Freshwater Fishes of Iran

Species Accounts

Cyprinidae

Introduction
with

Abramis
to Cyprinus

Revised:  03 June 2014
Back to Contents
 

Introduction

This family contains by far the most species in the Iranian freshwater ichthyofauna and is divided into two files Abramis to Cyprinus (here), and Garra to Vimba (see both in Contents).

The carp or minnow family is one of the most widespread and speciose families of fishes in the world, certainly the most speciose in fresh water and possibly the largest family of vertebrates (the Gobiidae may be the first). The family is found in North America, Eurasia and Africa. Other common names in English for species include barbels, breams, roaches, snow trouts, bitterlings, shiners, daces, chubs, barbs, "sharks", among many others. There are about 352 genera and over 2838 species (Nelson, 2006; Eschmeyer and Fong, 2011), about 8.9% of the world's fishes. In Iran, the family is represented by about ?32 native genera (interpretations of genera differ between authors) and at least ?73 species (with more to be described) found in all the major drainage basins. 

The minnow or carp family is comprised of small to very large fishes (1 cm and up to 3 m, with some of the largest members in Iran) characterised by throat or pharyngeal teeth in 1-3 rows, with a maximum of 8 teeth in a row, tooth counts and form are often characteristic of the genus or species, no jaw teeth, body form various from fusiform to compressed, lips are usually thin and not sucker-like (but can show hypertrophy), the upper jaw is bordered by the premaxillae bones and usually protrusible, barbels are absent or present in 1-3 pairs (not more than 2 pairs in Iranian species), body covered in cycloid scales, in some species easily lost, while the head is scaleless, no adipose fin, the anterior 4 vertebrae are modified for conduction of sound from the air bladder to the ear and are known as the Weberian apparatus, pelvic fins are abdominal in position, no pyloric caeca, air bladder usually present and well-developed, connected to the gut by a duct, and not enclosed in a bony capsule, no true stomach, branchiostegal rays always 3 in number, no true spines in the fins although in some the last unbranched dorsal fin ray (at the front of the fin) may be thickened and spine-like and in Cyprinus and Carassius the last unbranched anal ray is also thickened. The primitive chromosome number is 2n=50 but polyploidy is common and seen in Cyprinus, Carassius and in the schizothoracines. Collares-Pereira (1994) argues that the polyploid condition (e.g. 2n=100) is primitive or plesiomorphic.

There are 2-4 unbranched rays (including rudimentary ones) in the dorsal and anal fins followed by the more numerous branched rays (the last two branched rays are counted as one). The first pectoral and the first pelvic fin ray are unbranched and not included in counts. Pharyngeal teeth lie on a modified, fifth gill arch which can be seen or probed behind the shoulder girdle, just inside the gill opening. The arch has to be removed with dissecting equipment to count the teeth. Tooth counts are presented as a formula such as 2,5-4,1 which indicates 2 teeth in the outer left row and 4 on the inner right row. Teeth may be lost from major or minor rows so variant formulae are given after the principal one. A horny pad on the underside of the basioccipital bone of the skull is used to masticate the food against. Tooth form varies with the food - molar-shaped teeth are used to crush molluscs, flat but grooved surfaces for grinding plant food and sharp edged teeth for slicing various invertebrate foods.

Two subfamilies, the Alburninae and Leuciscinae, are paraphyletic but together seem to form a monophyletic group with a radiation about 20 million years ago, based on allozyme, cytochrome b, 16S rDNA and mitochondrial control region data from European cyprinids (Hänfling and Brandl, 2000; Gilles et al., 2001). These two subfamilies contain many Iranian genera. Zardoya and Doadrio (1999) analysed the cytochrome b nucleotide sequence of a variety of cyprinids, mostly European, and found support for two subfamilies Cyprininae (including barbins) and Leuciscinae (including cultrins, tincins, gobionins, phoxinins and alburnins + leuciscins). The origin of cyprinids is estimated at 38.9MYA and the separation of Cyprininae and Leuciscinae at 27.7MYA. They also found the phylogenetic utility of barbel possession to be limited as they were acquired independently in the two subfamilies. The number of rows of pharyngeal teeth were a more reliable phylogenetic marker, at least at the generic level. Perea et al. (2010) using mitochondrial and nuclear DNA give details of major cladogenetic events in the leuciscin lineages in the circum-Mediterranean, involving genera and species found in Iran.

Chen and Mayden (2009) investigated the major clades of cyprinids using a multiple nuclear gene approach and tentatively recommended elevation of certain subfamilies to family. This is not in general use at this writing and the Cyprinidae is retained as a single family here.

Durand et al. (2002) using cytochrome b DNA of Cyprinidae conclude that the the Middle East is an important interchange area for this freshwater ichthyofauna rather than a centre of speciation. The Middle East leuciscine cyprinids have Europe as an important Palearctic influence consistent with the Lago Mare dispersion while the the cyprinine cyprinids show three highly divergent lineages, namely one shared with the Euro-Mediterranean area (Barbus/Luciobarbus), a relict of the Lago Mare dispersion, one shared with Africa (Carasobarbus/Varicorhinus subgenus) and one with Asia (Garra). The Lago Mare dispersion occurred during a salinity crisis in the Mediterranean Sea 5.5 MY ago in the Late Miocene when freshwater fish were able to disperse through oligohaline or fresh water in the Paratethys Sea to reach the Middle East (Bianco, 1990). Some data of Durand et al. (2002) conflict with this scenario - the Carasobarbus clade that includes Barbus (= Tor) grypus shows a separation divergence later than the salinity crisis in the Pliocene when no migration route was available. But note that some authors place Barbus grypus in the Indian genus Tor and that evidently more work needs to be done on its relationships and on those of other species that have no evident Euro-Mediterranean relatives, but whose origins may well lie in the Oriental Region.

Other Middle Eastern cyprinid genera are regarded by Durand et al. (2002) as relicts of older colonization waves and show an eastern influence consistent with an Asian origin of the family Cyprinidae. Cyprinion has no sister species in the Euro-Mediterranean area and has been isolated in the Middle East since before the salinity crisis, 7.8-8.8 MY ago. Cyprinion may have entered the Middle East during the colonization event that isolated the genera Barbus sensu lato and Schizothorax in the European and Asian basins respectively. The divergence of these species is similar in time to the radiation of the Leuciscinae supposedly centred in Siberia based on fossil records. Siberia was probably an important dispersion centre for both Leucicinae and Cyprininae at that time. Otero (2001) describes a ?Barbus sp. (sic) from the Lower Miocene of Saudi Arabia showing an early date for the entry of cyprinids to the Afro-Arabian Plate.

Some species may enter brackish water but the family is primarily a freshwater one. Carps have extremely sensitive hearing via the Weberian apparatus and this is thought to account for their success. Carps produce an "alarm substance" when injured. This chemical stimulates other carps to flee and hide, another useful adaptation. Carps are remarkable for changes they undergo during the spawning season. Some fish, which are usually silvery, develop bright reds and yellows. Nuptial, pearl or breeding tubercles develop on the head, scales and fin rays often in distinct patterns, and there are swellings of the head or fin rays in some species. These changes are most apparent in males. Tubercles and swollen rays are used to clasp females during the spawning act. Generally males have longer pectoral fins than females. Tubercles are also used to fight other males and defend and clean nests. Colour attracts females for mating. Nest building males are larger than females, the reverse of the situation in most fishes where egg-bearing females are the largest. Not all species build nests and some simply broadcast eggs over weed, gravel or sand. Fractional spawning is common in carps. This is a prolonged spawning season which ensures no single batch of eggs is lost to unfavourable, temporary environmental changes such as floods. Carps are mostly omnivores, feeding on small crustaceans, insects and some minute plants but some specialise in eating large plants, or other fishes. Diet is reflected in pharyngeal tooth shape as mentioned above. Gut length is important too. A long intestine indicates a reliance on plant material which takes longer to digest. A simple, s-shaped gut is found in insectivorous fish. A black peritoneum is thought to protect gut bacteria from damaging light. The bacteria aid in breaking down the strong cell walls of plants. Size and shape of the mouth are also indicative of diet. Carps are found in many diverse habitats from swift, cold streams to warm bogs. These are schooling fishes, especially when young.

Carps play an important role in fresh waters as food for other fishes and some species are commercially important as bait fish, as sport fish or as food in Asian countries. Raising minnows as bait and as forage fish for sport fish is a big business in the U.S.A. They are an important element in the commercial aquarium trade and certain species are used in experimental studies by scientists. Cyprinids were also important in the past, sacred fish ponds being reported from Mesopotamia in 3000 B.C., and in Iran today cyprinids associated with mosques and shrines are "sacred". A general review of Eurasian cyprinids is given by Bănărescu and Coad (1991).

Carp family members are particularly important in Iran in aquaculture. The "Chinese carps" (Cyprinus carpio or common carp, also native to Iran, Ctenopharyngodon idella or grass carp, Hypophthalmichthys molitrix or silver carp, and to a lesser extent Hypophthalmichthys nobilis or bighead carp) are the main species used in warmwater culture in almost all the provinces of Iran. Common, grass and silver carps are processed into fish fingers in Iran (Iranian Fisheries Research Organization Newsletter, 25:1, 2000). Danesh-e-Khoshashi (1998) describes facilities and methods used for spawning Chinese carps in Gilan Province. The production of Chinese carp fingerlings has been relinquished to the private sector in Iran. The silver carp catch increased from none in 1989 to 24,720 t in 1994 (Food and Agriculture Organization, Fisheries Department, 1996). Chinese carp production peaked in 2006 at more than 77000 t according to Salehi (2009) who also reviews carp farming costs. Chinese carp fingerling production was 22.7 million in 1996 (Bartley and Rana, 1998a). Stakei (1999) studied nutrients, BOD and COD in manured polyculture ponds with Chinese carps. A review of world cyprinid culture, with special reference to the Chinese carps, is given by Billard (1995).

Rana and Bartley (1998a) give details of carp aquaculture in Iran. They note that silver carp production increased 11% per year between 1991 and 1996 and bighead carp 7%. Most carp production occurs in the provinces of Gilan, Mazandaran and Khuzestan and is a private sector enterprise. Carp broodstock is selected based on head size, colour and gill structure (surface and shape). Adults are replaced after 3-4 years. Circular concrete tanks are used for spawning and have egg collecting and incubation devices which reduce handling to the minimum. The young carp are grown to market size in ponds or complex fish farms. In 1994, there were 2583 registered farms with a water surface area of about 8000 ha. Organic and inorganic fertilizers are used along with supplementary foods. Fertilizers include urea (135-1500 kg/ha/yr), ammonium phosphate (80-575 kg/ha/yr) and manure (3-10 tonnes/ha/yr). Supplementary diets include a variety of grains (100-6000 kg/ha/yr) or, for intensive monoculture of common carp, high protein pellets (30-40%). Fingerlings are stocked in March-April at a density of 2000-6000 per hectare and sold between November and February. Production is 1.6-5.5 tonnes/ha. Cultivated carps are susceptible to fungal infections as detailed by Ebrahimzadeh et al. (2000) for the Safid River Fish Farm Centre where 31 species of fungi were isolated and Firouzbakhsh et al. (2005) where 39 fungal species were identified from gill lesions in common, silver and grass carp on five fish farms in Mazandaran.

Rice fields in Iran are now being considered for fish culture. Experimental production of 300-500 kg per hectare of "carp seed" (presumably young fish) an 750-1000 kg of fish and ducks in the autumn after the paddy is harvested (Iranian Fisheries Research Organization Newsletter, 22:2, 2000). In the early 1970s intensive carp culture yielded only half the profits of rice culture (Carl Bond archives, Oregon State University, Corvallis).

Experiments in the Caspian region for artificial propagation of Aspius aspius and Barbus (= Luciobarbus) brachycephalus to enhance stocks and for farming Rutilus frisii and Abramis brama using mono- and polyculture along with Chinese carps have been carried out (Iranian Fisheries Research and Training Organization Annual Report, 1992-93; Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 77-78, 1995). There are about 3000 fish farms producing over 98% of the cultured fish in the country. Yearly production of all cultured fish has increased from 4753 tonnes in 1985 to 45,134 t in 1990. Production of carps in government hatcheries has risen as follows: 2.19 million fingerlings in 1983, 5.04 million in 1984, 12.84 million in 1985, 20.83 million in 1986, 19.05 million in 1987, 50.00 million in 1988, 50.80 million in 1989, 97.70 million in 1990, 58.00 million in 1991, and 50.00 million in 1992. In addition private sector production probably equals these figures (Emadi, 1993a). Polyculture of common, bighead and silver carp has been tried in Iran (Kamaly, 1991). Fish were stocked in four 200 sq m ponds at three densities in polyculture (2700, 3750 and 4750 by species) and at one density in monoculture (9500) fish per hectare. Bighead and silver carp attained a mean weight of 526 and 498 g in polyculture and common carp averaged 343, 190 and 100 g in the same culture but only 13.6 g in monoculture. The growth rate in summer averaged 94.4, 93.7 and 76.1% for silver, bighead and common carp in polyculture and 71.9% for common carp in monoculture. Pen culture in the Caspian Sea has been investigated for Cyprinus carpio and the various Chinese carps (Iranian Fisheries Research and Training Organization Annual Report, Tehran, 1992-93). Semi-artificial breeding of grass, silver and bighead carps has been carried out in Iran (Iranian Fisheries Research and Training Organization Newsletter, 6:3-4, 1994; Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39, 1996). Hormone injections were used to induce breeding of fish held in a round trough for spawning with a rectangular egg collection trough and a round egg hatching trough. Spawning occurred within 6.5-12.5 hours of injection. The percentage of hatched larvae in this semi-artificial method was higher than a control artificial method where eggs are kept in incubators. The increase was 6% for grass carp, 33.72% for silver carp and 16.7% for bighead. Active larvae increased from 180,000 to 450-500,000 for grass carp, from 157,000 to 400-450,000 for silver carp and from 680,000 to 970,000 for bighead carp. Additionally female breeder mortality was 3.37% less for grass carp and 45.19% less for silver carp.

Many carp species can be caught on hook and line by various angling techniques but outside the larger rivers of Khuzestan and the Caspian shore this hobby is not much pursued. Even small species and specimens can give some sport on light tackle such as worm baited hooks including Luciobarbus barbulus, Carasobarbus luteus, Alburnus mossulensis, Cyprinion macrostomum and Garra rufa among others.

Fingerlings of Labeo rohita, an Indian carp, were imported to Gilan in Iran in 2004 to enrich the diversity of cultured fish and increase protein production. There is always the potential for escapes and establishment of this exotic. Additionally, Mortezavi Zadeh et al. (2012) report on pond-cultured Catla catla, Labeo rohita and Cirrhinus reba (Indian carps) in Khuzestan where the climate may well favour any escapees - the fish survived a January-September  temperature regime of 7-34°C. There is also an extensive aquarium fish culture industry in southern Iraq, an additional source of exotic cyprinids (Coad, 2010).
 


Genus Abramis
Cuvier, 1816 
 

The bream genus comprises 4 species found in Europe, Asia Minor and the Caspian and Aral Sea basins. There are 2 species in Iran but see also Blicca and Vimba.

The genus is characterised by a strongly compressed and deep body, a scaleless keel between the vent and pelvic fins, a scaleless groove on the back in front of the dorsal fin but not behind the fin, pharyngeal teeth in 1 row, compressed and with a groove on the grinding surface, dorsal fin short and spineless, anal fin long to very long, and lateral line decurved.

Durand et al. (2002) studying cytochrome b data concluded that this genus is not monophyletic since A. ballerus and A. sapa are placed basal to a group of species including A. brama, Blicca bjoerkna, Vimba species, Acanthalburnus microlepis and Acanthobrama.

.Abramis brama
(Linnaeus, 1758)


Anzali Shore, June 2012, courtesy of K. Abbasi

Abramis brama, aquarium fish, from Wikimedia Commons
Abramis brama, aquarium fish, from Wikimedia Commons

Common names

سيم (sim or seam = silver), ماهي سيم (= mahi-ye sim, meaning silver fish).

[capag, chakag, chapakh or chipakh, all in Azerbaijan; gundogar tarany (topi) in Turkmenian; vostochnyi leshch or Oriental bream in Russian; common, bronze, eastern or carp bream].

Systematics

Cyprinus Brama was originally described from Sweden.

Abramis brama orientalis Berg, 1949 is reported for the Caspian and Aral Sea basins but Koshara and Izyumov (1991) restricted this subspecies to the Aral Sea with the type subspecies in the Caspian Sea basin. They did not examine any Iranian material. Kozhara and Mironovskiy (1988) using numbers of pores in the seismosensory canals for samples taken over a wide range of this species identified 8 population groups but did not recognise subspecies. Some earlier works also indicate that no subspecies exist (see Reshetnikov et al., 1997).

Caspian material reportedly has more gill rakers, fewer vertebrae and fewer scales than the type subspecies from the Baltic Sea basin (Berg, 1948-1949) but further study over the whole range of the species is needed to clarify the situation, analyzing for clines. The Iranian populations are referred to the type subspecies for the moment. The type locality of this subspecies is the Aral Sea at Muinak and Lake Yaskhan in the Uzboi.

Khara et al. (2007; 2007) compared fish from the Anzali wetland and the Caspian Sea, and the Caspian Sea and Aras Dam, both meristically and morphometrically. Significant differences were noted in particular for morphometric characters in the former comparison and morphometrically and meristically in the second. These differences were attributed to differing habitats and environmental conditions. Ghasemi et al. (2007) used 5 microsatellite loci in comparing Iranian and Azeri bream and found Iranian stocks have reduced genetic variability attributed to inbreeding and genetic drift. Khara et al. (2009) compared fish from the Anzali Wetland the southern coast of the Caspian Sea in Iran and the southwest coast in Azerbaijan using mtDNA. The greatest genetic diversity was found in Azerbaijan which was significantly different from the Iranian samples, which were not themselves significantly different.

Abramis brama bergi Grib and Vernidub, 1935 (preoccupied by Abramis sapa bergi Belyaeff, 1930 according to Eschmeyer et al. (1996)) was originally described from the Aral Sea at Muinak and is also found in the Uzboi Valley of Turkmenistan, north of the Iranian border (Berg, 1948-1949). It was replaced by Abramis brama orientalis.

A syntype of Cyprinus brama is in the Natural History Museum, London as a skin under BM(NH) 1853.11.12:147 (Eschmeyer et al., 1996).

Artificial hybrids with Rutilus frisii kutum and Rutilus rutilus (may involve R. caspicus) have been bred in Iran (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 39-40, 1996).

Key characters

The scaleless keel on the belly, deep body, high number of branched rays in the anal fin (22-30), modally 9 branched dorsal fin rays, and uniserial pharyngeal teeth are key characters.

Morphology

The mouth is small but highly protrusible. There is a strong dorsal ridge anterior to the dorsal fin. Dorsal fin with 3 unbranched and 8-10, usually 9, branched rays, anal fin with 3 unbranched and 22-30 branched rays. Lateral line scales 48-60. The lateral line is moderately decurved. Scales are regularly arranged, sheathing the anal fin base. Scales have numerous fine circuli but only relatively few posterior and even fewer anterior radii. In a fish about 6 cm long there are as few as 8 total radii. The focus is almost central and the anterior scale margin is wavy. There is a pelvic axillary scale. The ventral keel between the pelvic fin bases and the anal fin is well-developed. Gill rakers number 18-30 and are short, reaching the raker below when appressed. They are strongly tuberculate on the inner surface. Vertebrae 38-47, usually 42-44 in the Caspian populations (lower counts in literature may not include 4 Weberian vertebrae). The chromosome number is 2n=50-52 (Klinkhardt et al., 1995). The chromosome number based on fish from the Iranian coast of the Caspian Sea is 2n = 50 with the number of arms NF = 82 and the karyotype being 8 pairs of metacentric, 8 pairs of submetacentric and 9 pairs of acrocentric chromosomes (Nahavandi et al., 2001).

Pharyngeal tooth formula modally 5-5, with variants of 6-5 (2.2-4.8%), 5-4 (2.2-4.4%) and 4-5 (8.6%) for collections from the Caspian and Aral seas basins in former Soviet waters (Vasil'yeva and Ustarbekov, 1991). Other variants are summarised in Tadajewska (1998). Teeth bear a small hook at the tip in the main row and have long, narrow and flat crowns. In young fish, the hook is more pronounced and the crown has a few tubercles or a series of serrations. The gut is s-shaped with a small anterior loop.

Khar et al. (2007) compared this species from the Caspian Sea and the Anzali wetland and found significant morphometric, but not meristic, differences, attributing this to habitat conditions.

Meristic values for Iranian specimens are:- dorsal fin branched rays 9(12) or 10(1); anal fin branched rays 24(3), 25(3), 26(1), 27(4) or 28(2); pectoral fin branched rays 16(7) or 17(6); pelvic fin branched rays 8(13); lateral line scales 49(2), 50(2), 51(3), 52(3), 54(1) or 55(2); total gill rakers 23(3), 24(2), 25(6), 26(1), or 27(1); total vertebrae 44(12) or 45(1); and pharyngeal teeth 5-5(12).

Sexual dimorphism

Males bear tubercles on the head, body and fins. Scale tubercles appear singly, in pairs or occasionally as 3 per scale. There is some evidence of differences in gill raker counts between the sexes but sometimes the males have higher mean counts and sometimes the females. Abdurakhmanov (1962) reports eye diameter, greatest body depth and predorsal distance to be greater in females and dorsal fin base length, pectoral and pelvic fin lengths and interorbital width to be greater in males from Azerbaijan.

Colour

In Dagestan, the resident form is darker in colour than the semi-anadromous form (Shikhshabekov, 1969). Overall colour is silvery. The iris is silvery with a little grey pigment on the upper part. The dorsal and caudal fins are pale grey, almost transparent, to a greyish-blue, the pectoral fins may be grey or colourless, and pelvic fins are colourless. All fins except the pectorals have black tips. Large fish are a dark olive-green on the back and bronze on the flanks and old fish may have all fins black. The peritoneum is silvery to light brown in preserved fish.

Size

Attains 90.0 cm total length and 11.55 kg, possibly 1.0 m and 16.4 kg (Machacek (1983-2012), downloaded 27 July 2012).

Distribution

Found from the British Isles across Europe north of the Pyrenees and Alps eastwards to the Black, Caspian and Aral sea basins although not in western Transcaucasia. In Iran it is found from the Astara to the Atrak rivers in the whole Caspian Sea basin (Kozhin, 1957) including the Anzali Mordab, its outlets, tributaries and shore, and the Siah-Keshim Protected Region (Holčík and Oláh, 1992; Riazi, 1996; Kiabi et al., 1999), the Safid River (Abbasi et al., 1999), Gorgan Bay (Derzhavin, 1934), and freshened areas of the Caspian Sea. It is also found in the Aras Dam (Khara et al. 2007).

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin where it has been reported by Aliev et al. (1988).

Zoogeography

This species is part of a northern European and northern Southwest Asian fauna whose zoogeographical history has not been researched. It origins may lie in a Danubian or Sarmatian fauna.

Habitat

The bream prefers still water and is low in numbers even in rivers with weak current. Abundant littoral vegetation and a very muddy bottom are favoured in lakes for reproduction and feeding respectively. It retreats to deeper water in winter, forming schools numbering in the many thousands, packed densely together (Muus and Dahlstrøm, 1999). It is more numerous in the Anzali Mordab along the Caspian coast of Iran (Jolodar and Abdoli, 2004).

This species can tolerate high temperatures of 33-34°C in southern areas like Iran for a time but above 28°C growth rate decreases. Adults can live in a salinity of 12.9‰, perhaps 14‰, and eggs may be fertilised at a salinity of 10.2‰. However preferred levels are 2-4‰. Salinity and water level changes have significant effects on abundance in this species. Population densities vary markedly in both fresh and brackish water populations.

Bream living in the Caspian Sea basin are semi-migratory. They feed in the brackish sea but spawn and winter in the lower reaches and deltas of large rivers. A spring migration up rivers begins with ice melt or warmer temperatures in the sea and after spawning the fish return to disperse and feed in the sea. In the fall the fish migrate into the deeper parts of river deltas. In Russian parts of the Caspian they are found at depths not exceeding 4-5 m but Knipovich (1921) reports them at 14.6-16.5 m, possibly deeper, in the Iranian Caspian Sea.

There were spring and winter migrants in the southwestern Caspian including the Anzali Mordab (A. M. Shukolyukov in Berg, 1948-1949). The spring bream had a longer snout, deeper head, lower body, lower dorsal and anal fins, and more scales. The spring bream entered the Mordab for spawning only while the winter bream overwintered in bottom pools. Changing conditions in the mordab environment in the late 1980s and the 1990s may have altered this migration. Riazi (1996) reports that this species migrates into the Siah-Keshim Protected Region of the Anzali Mordab.

Age and growth

The resident form in Dagestan is slightly inferior in length (2-3 cm), weight and age to the semi-anadromous form (Shikhshabekov, 1969). In Dagestan, the resident form becomes sexually mature at 3 years for females and 2 years for males at lengths of 23-26 cm and weights of 200-240 g while the semi-anadromous form matures at 4 years and a length of at least 25-28 cm and a weight of 250-300 g. In Uzbekistan females mature at lengths ranging from 10.5 to 27 cm in different reservoirs, usually at age 3 (Kamilov, 1994). Maturity is attained at a younger age in southern waters generally in this species and this probably applies in Iran. The maturity range is 2-10 years with males often maturing a year earlier than females. Females predominate in the older age groups.

Maximum age exceeds 32 years although in southern waters the maximum age does not exceed 15 years. Semi-migratory bream of the Caspian Sea have a fast growth rate and a short life cycle, reaching 37.5 cm standard length by age 8.

Most fish examined by Razivi et al. (1972) from commercial catches in Iran were 3-6 years old, 25.6-39.8 cm long and weighed 249-950 g. Over a three year period there was a decline in average age. Young and immature fish formed most of the catch in 1998-1999 when one-year-old fish comprised 20.3% and two-year-old fish 37.3%. The average length, weight and age for 1998-1999 were 22.5 cm, 212.2 g and 2.4 years. The rate of recruitment was 4.6% in 1991 and 2.7% in 1992 (Saiad Borani, 2001). Abdolmalaki (2005b) studied Caspian Sea fish from Iran and found mean fork length, weight and age to be 21.7 cm, 191 g and 2.72 years, respectively. The length-weight relationship was W = 0.2312L2.9 and von Bertalanffy growth parameters were Lt = 45[1-exp-0.125(t  2.768)], and the instantaneous rate of total (Z), natural (M) and fishing mortality (F) were 0.92 year-1, 0.28 year-1 and 0.64 year-1, respectively. The exploitation rate (E) was 0.7. Biomass was calculated as 46.362 t and the maximum sustainable yield was 14.99 t.

Food

Young fish feed on zooplankton. Adults use a strong sucking power and a tube-like snout to feed on invertebrates and detritus in mud. This sucking action leaves evident "bream pits" in soft mud, depressions about 10 cm across. In the northern Caspian Sea food items include Cumacea, Corophiidae, the clams Adacna (69% by weight) and Monodacna, Tendipedidae (= Chironomidae), Polychaeta, Gammaridae, Mysidae, and Oligochaeta. When overcrowded or in turbid conditions, plankton may be eaten in addition to the normal foods (Muus and Dahlstrøm, 1999). Large specimens may feed on small fishes. A specimen from the Langarud, Gilan, 158.6 mm standard length, contained chironomids.

Reproduction

Bream enter the Kura River from December to February with a peak in January (Berg, 1959) and travel some distance upriver. These fish have an average length of 31.1 cm and an average weight of 633 g. Length and weight in Azerbaijan vary from 25.4 to 31.9 cm and 306 to 681 g. Bream enter the Anzali Mordab, the main spawning area in the southern Caspian, in the first half of March until the beginning of May. Males precede females on the spawning ground by about 3 days and males outnumber females by about 3 to 1. Spawning begins in the first half of April in shallow water and lasts until mid-May. Fecundity in Dagestan reservoirs reaches 191,000 eggs (Shikhshabekov, 1969), in Uzbekistan reservoirs 772,000 eggs (Kamilov, 1994) and a maximum elsewhere of 941,000 yellowish eggs is reported. Bream spawn repeatedly with different partners and although most bream spawn only once a year, multiple spawnings are known. Spawning occurs in masses over a period of 2-3 days triggered by temperatures of 12-13°C or above. The commonest spawning temperature for the species overall is 16-18°C. Spawning is most intensive at night in some populations while others show late morning and late afternoon peaks. There is much splashing of the water by their tails and the noise can be heard some distance away although the fish are easily scared into deeper water by any noise like human voices. Males probably defend territories, attracting females and scaring other males away. There can be up to 2.3 million eggs per sq m however, suggesting that many fish may spawn in the same area. Eggs are deposited in quiet water, most commonly at depths of 20-80 cm, and they adhere to aquatic plants or flooded land plants. Eggs are up to 1.9 mm in diameter.

Parasites and predators

Jalali and Molnár (1990a) record the monogenean Dactylogyrus zandti from this species in the Safid River. Sattari and Faramarzi (1997) record Caryophyllaeus fimbriceps from 28% of bream in the Anzali lagoon. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the monogenean trematodes Dactylogyrus zandti and D. wonderi. Masoumian et al. (2005) report the protozoan parasites Ichthyophthirius multifilis and Trichodina perforatafrom this species in the Aras Dam in West Azarbayjan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. elegans from fish in the Safid River. Sattari et al. (2004, 2005) survey this species in the Anzali wetland, recording Raphidascaris acus larvae. Pazooki et al. (2007) record various parasites from localities in West Azarbayjan Province, namely Ligula intestinalis, Digrama sp., Argulus foliaceus and Caryophyllaeus laticeps. Sattari et al. (2007) record the cestode Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus and Gyrodactylus sp. in this species in the Anzali wetland of the Caspian shore and also mention that the monogenean Diplozoon sp. is also known from this species in the Iranian Caspian Sea. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species.

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984). Various predatory fishes take bream including Huso huso, Perca fluviatilis, Sander lucioperca, Aspius aspius and Silurus glanis but this is comparatively rare especially when bream exceed 20 cm in length. Birds such as grebes, herons, divers and cormorants are also predators.

Economic importance

This species is an important food fish being both tasty and of large size. In addition it can live out of water for some time and thus remain fresh while being transported to market.

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Anzali region from 1901-1902 to 1913-1914 the catch was 2283 to 419,117 individuals, for the Safid River region from 1908-1909 to 1917-1918 the catch was 17,195 to 474,200 individuals (rising steadily but falling in 1917-1918) with no fish reported in the years 1899-1900 to 1907-1908 and in 1918-1919, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 20,600 to 1,381,500 individuals with no clear trend, the catches varying markedly from year to year. The commercial catch in Iran from 1956/1957 to 1961/1962 varied from 0 to 158 kg (Vladykov, 1964), from 1965/66 to 1968/69 varied from 0 to 29 tonnes (Andersskog, 1970) and from 1963 to 1967 from 0.5 to 16.0 tonnes (with no reported catch in the first 3 years)(RaLonde and Walczak, 1970b). The catch in the Bandar-e Anzali region from 1933/34 to 1961/62 varied between only 2 kg and over 1394 t with some years reporting no catches. Holčík and Oláh (1992) report a catch of 34 kg in the Anzali Mordab for 1990 and for the period 1932-1964 catches ranged from none to 1133.5 tonnes annually. The total catch of the Northern Shilot (Fisheries Company) from 1965/66 to 1968/69 varied between 13 and 74 t (RaLonde and Walczak, 1972). There are obviously wide variations in annual catches and/or in reporting statistics. The general trend is one of decline in catches with large fish being caught and the average stock size being lowered, resulting in a decreased spawning success. This species has a deep body and immature fish are easily caught. The catch in the Anzali Mordab was important until the end of the 1940s but had virtually disappeared by the 1980s (Petr, 1987). Abdolmalaki (2005b) gives a total catch of 17 t for the 2000-2001 fishing season, only 0.1% of the commercial catch in Iranian coastal waters of the Caspian Sea. In contrast, the total catch for Iranian waters was estimated at 26.3 tons of which 15.4 tons was from beach seines; most fish were immature and undersized (Abdolmalaki, 2006a).

In former Soviet waters of the Caspian Sea, the age composition in commercial catches was 2-10 years, with the great majority being 3-5 years old. Trawls, seines, pound nets and gill nets are used in the northern Caspian Sea to catch the bream with 60-70% being taken in spring. Spawning and breeding farms were established in the former Soviet Union to rear young fish. Catches in the Volga-Caspian and Ural regions has been as high as 344,900 centners, prior to 1930, and in the Aral Sea in 1931 the catch was 115,200 centners.

Mono- and poly-culture of this species has been carried out in Iran (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 77-78, 1995). Polyculture comprised 70% Abramis brama, 20% silver carp (Hypophthalmichthys molitrix) and 10% grass carp (Ctenopharyngodon idella) and gave a greater yield than monoculture. From an average initial weight of 30 g, fish attained averages of 188 or 211 g in monoculture (average 200 g) and 221 or 278 g (average 250 g) in polyculture with maximum weights of 300 or 580 g at the end of two one-year periods. Water temperatures were 9-33°C (Annual Report, 1994-1995, Iranian Fisheries Research and Training Organization, Tehran, p. 38-39, 1996; Danesh-e-Khoshashi, 1997).

A state supported stocking programme has released about 70-80 million fingerlings into the Anzali Mordab, all descended from a single pair mating 8 years ago (Rana and Bartley, 1998a; 1998b). These fish are intolerant of low oxygen and so perform poorly under pond conditions. Stocks may be imported from Azerbaijan in the future (Rana and Bartley, 1998b). The release of 70.46 million fry in 1992-1993 to 1998-1999 period has not been successful in restoring the stocks in Iran. Stock depletion was attributed to improper fishing methods, pollution, destruction of spawning grounds, presence of predatory Esox lucius and Silurus glanis in fry stocking areas, and lack of necessary arrangements in regard to artificial spawning (Saiad Borani, 2001).

The roe or eggs of this species have been implicated in poisoning (Halstead, 1967-1970; Coad, 1979b) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death.

This species has been used in Iran for experimental studies, e.g. on the toxicity and LC50 of phenol and 1-naphthol (Shariati et al., 2004).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and in aquaculture.

Conservation

The subspecies has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985).

RaLonde and Walczak (1970b) reported that 90% of the bream caught in Iran in 1970 were immature and the stock was in danger of extinction. About 19-20% of commercial catches in the Volga region are from hatchery raised stock (Petr, 1987) and it was thought that stocking could help this species in Iran. During the 1980s and 1990s there were practically no catch figures for this species in Iran. Artificial propagation began in 1986 on an experimental basis and 6 million fish were released (Ghenaat Parast, 1993). In 1992-1993 (an Iranian calendar year), 2.4 million fingerlings were released into the Anzali Mordab and nearby rivers, a 100% increase over the previous year (Abzeeyan, Tehran, 4(2):VI, 1993). Total production in government hatcheries for 1990 was 0.66 million fingerlings, in 1991 2.28 million and in 1992 5.3 million fingerlings (Emadi, 1993a). Fingerling production was 11.217 million in 1995 and 8.5 million in 1996 (Bartley and Rana, 1998a; 1998b). In 1999-2000, 20 million juveniles were released (Iranian Fisheries Research Organization Newsletter Newsletter, 23:4, 2000). From October to March 2000, 14 million juveniles raised in the Shahid Ansari aquaculture and breeding centre in Gilan were released into the Caspian Sea and neighbouring water bodies (Iranian Fisheries Research Organization Newsletter, 26:2, 2001). Illegal fishing and non-standard nets threaten the stocks (Annual Report, 1995-1996, Iranian Fisheries Research and Training Organization, Tehran, p. 19-20, 1997). Billard and Cosson (2002) give an annual production of 15 million alevins.

Ramin (1997) details studies on the artificial breeding of this species in Iran, based on 38 brooders, with the goal of saving it from extinction. Gonadotropic hormone extracted from the pituitary of the common carp was used to induce brooders. One or two doses at 5-6 mg/kg body weight gave optimum stripping of eggs at 18°C. Fertilisation rate was 75-95% and hatching rate was 75-85%. Incubation took nearly 4 days at 18-21°C. The grey, pink or yellow eggs numbered 9142-60,050 per spawner with a swelled diameter of 1.0-1.2 mm. The yolk sac was absorbed after 72 hours and newly hatched larvae were 2.9-3.7 mm long.

Khara et al. (2009) (see above) carried out their molecular study in order to determine sources for broodstock to increase genetic diversity after losses from overfishing, pollution and loss of spawning regions. Koohilai et al. (2010) studied optimum doses of various hormones used to stimulate ovulation.

Kiabi et al. (1999) consider this species to be vulnerable in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, few in number, habitat destruction, limited range (less than 25% of water bodies), not present in other water bodies in Iran, and present outside the Caspian sea basin. Nezami et al. (2000) consider this species to be endangered because of overfishing, habitat destruction and spawning grounds degradation. Shapoori and Gholami (2005) found the fishery not to have changed despite millions of  fry being stocked.

Further work

Stocks should be carefully monitored on a continuing basis and efforts made to resurrect this commercial species.

Sources

The chief literature summary for earlier works is Backiel and Zawisza (1968) although little apparently refers to the Caspian basin populations and even less to those of the Iranian shore. Nevertheless this work gives a general overview of biology and general comments above are based on it.

Iranian material: CMNFI 1970-0542, 4, 75.4-173.7 mm standard length, Gilan, Old Safid River estuary (37°23'N, 50°11'E); CMNFI 1970-0543A, 1, 70.0 mm standard length, Gilan, Caspian Sea at Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1971-0343, 1, 158.6 mm standard length, Gilan, Langarud at Chamkhaleh (37°13'N, 50°16'E); CMNFI 1980-0127, 3, 166.1-170.1 mm standard length, Gilan, Caspian Sea near Hasan Kiadeh (37°24'N, 49°58'E); CMNFI 1980-0142, 1, 160.6 mm standard length, Gilan, Nahang Roga River (no other locality data); CMNFI 1980-0906, 3, 105.6-176.0 mm standard length, Gilan, Caspian Sea basin (no other locality data).

Abramis sapa
(Pallas, 1814)

Common names

سابا, سبا (saba, from the species name), سيم (sim = silver), ماهي سيم كند پوزه (= mahi sim kondpuzeh, meaning bluntsnout silver fish).

[pori or poru, both in Azerbaijan; tarashka, taran' and rybets, erroneously in Azerbaijan; yuzhnokaspiiskaya beloglazka or South Caspian white-eye bream in Russian; white-eye bream, southern white-eye bream, Danube bream].

Systematics

Cyprinus Sapa was originally described from the Sura, Samara and Kinel' rivers in the Volga River basin. No types known.

May be placed in the genus Ballerus Heckel, 1843 (see Hensel (1978), Shcherbukha (1973), Howes (1981), Bogutskaya (1986) and Bogutskaya and Naseka (2004) for various opinions). The nominate subspecies was described from the Volga River and tributaries.

The subspecies reported from the southern Caspian Sea basin is Abramis sapa bergi Belyaev, 1929, described from the Kura River in Azerbaijan. Eschmeyer et al. (1996) date this subspecies to 1930 although the article is dated 1929. Recognition of subspecies is disputable (Reshetnikov et al., 1997).

Key characters

The scaleless keel on the belly, deep body, very high number of branched rays in the anal fin (31-44), modally 8 branched dorsal fin rays, and uniserial pharyngeal teeth are key characters.

Morphology

Dorsal fin with 2-3, usually 3, unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 31-44, mostly 34-38 branched rays, pectoral fin branched rays about 15 and pelvic fin branched rays about 8. Lateral line scales 42-55, mostly 51-52, regularly arranged over the body. Scales bear numerous very fine circuli, an almost central focus, numerous to few posterior radii (quite variable between scales of similar size) and few to none anterior radii. The anterior scale margin is wavy. A pelvic axillary scale is present. There is an evident, scaleless keel on the belly between the pelvic fin bases and the anal fin. Gill rakers 18-25, short, reaching the raker below or almost the second raker when appressed. Vertebrae 45-48. Pharyngeal teeth 5-5, with elongate, narrow and flattened, concave or rounded crowns below a hooked tip. The gut is s-shaped with a small anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Belyaev (1929) for Kura River fish gives lateral line scale counts as 48(3), 49(6), 50(24), 51(50), 52(54), 53(16) or 54(7) and anal fin branched rays as 32(1), 33(5), 34(22), 35(32), 36(38), 37(47), 38(25), 39(9), 40(5), 41(2) or 42(1). This subspecies is distinguished from the type form in the Black Sea (Don River) by fewer lateral line scales and anal fin branched rays, a longer snout, smaller eyes, less deep body, lower dorsal fin, shorter anal fin, and longer postorbital length.

Sexual dimorphism

Unknown.

Colour

The Caspian subspecies has a dark back with a bluish tint, flanks and belly are silvery, fins are a greyish-white and sometimes have a black margin, and the iris is silvery. The peritoneum is dark brown in preserved fish.

Size

Attains 41 cm and 0.8 kg.

Distribution

Found in the basins of the Black, Caspian and Aral seas. Reported from the gut of a Silurus glanis in the Anzali Mordab (Derzhavin, 1934) but not found in recent years (Holčík and Oláh, 1992). Other reports are from the lower Safid River at Hasan Kiadeh (Belyaev, 1929; Derzhavin, 1934) and in the Aras River at Karadonly (Berg, 1948-1949).

Zoogeography

This species is part of a northern European and northern Southwest Asian fauna whose zoogeographical history has not been researched.

Habitat

This species feeds in brackish water but spawns and overwinters in the lower reaches of rivers. It is commonest along the western shore of the middle and southern Caspian Sea.

Age and growth

Females are 28-29 cm long on average, maximum 39 cm, while males are about 24 cm, maximum 30 cm (Belyaev, 1929). Males and females mature at 2-3 years and life span is 5 years in Azerbaijan (Abdurakhmanov, 1962).

Food

Food items include small molluscs, crustaceans and insect larvae as well as some plant fragments and detritus. Young feed on zooplankton.

Reproduction

A migration into rivers, particularly the Kura, occurs in winter when temperatures fluctuate from 5 to 10°C (Belyaev, 1929). The run begins in November and peaks in January. The Kura migration was once over 700 km from the mouth. Spawning occurs in rivers with gravel bottoms or dense vegetation from April to May. Fecundity reaches about 150,000 eggs with diameters up to 1.8 mm. Eggs adhere to stones or plants.

Parasites and predators

Eaten by Silurus glanis (Derzhavin, 1934).

Economic importance

Up to 1-2 million fish were caught in the Kura at spawning (Belyaev, 1929). The annual average catch in Azerbaijan in 1931-1935 was 1,860,000 fish weighing 6200 centners.

Conservation

The subspecies A. sapa bergi has been proposed for inclusion in the "Red Book of the U.S.S.R." which forms the basis for measures to protect species (Pavlov et al., 1985). It has always been very rare in Iran and its absence from the Anzali Mordab may be due to loss of spawning grounds (Holčík and Oláh, 1992). Lelek (1987) considers this species to be rare to vulnerable in Europe.

Further work

The status of this species in Iran should be assessed by field surveys. It is apparently quite rare and was not caught during two collecting trips along the Caspian shore in the 1970s. It is recorded only from two localities in Iran in 1929 and 1934.

Sources

Iranian material:- None available, based on literature reports.

Comparative material:- CMNFI 1986-0458, 2, 209.0-211.7 mm standard length, Germany, Danube River (48º58'N, 12º18'E); BC 59-301, 2, 136.3-154.2 mm standard length, Ukraine, Tisa, Danube drainage (no other locality data).

Genus Acanthalburnus
Berg, 1916

This genus contains only 2 species, both found in Iran. Berg (1948-1949) characterises it as similar to Alburnoides but with the last unbranched dorsal fin ray thickened into a spine which is strong basally but becomes thinner and flexible on about the last third of the ray length. Pharyngeal teeth are in 2 rows as opposed to 1 row in Abramis. Durand et al. (2002) include this genus in the Abramis clade based on cytochrome b data while Perea et al. (2010) using mitochondrial and nuclear DNA propose synonymy with Acanthobrama.

Acanthalburnus microlepis
(De Filippi, 1863)

Safid River, August 2010, courtesy of K. Abbasi
Safid River, August 2010, courtesy of K. Abbasi

Common names

مرواريد ماهي لب نازك (= morvaridmahi-e labnazok), kuli.

[garagas or taxta balig, both in Azerbaijan; chernobrovka and napota in Russian; blackbrow bleak].

Systematics

Abramis microlepis was described from the "Kur, presso Tiflis" (= Kura River near Tbilisi, Georgia) and the holotype is in the Istituto e Museo di Zoologia della R. Università di Torino under MZUT N.673 (Tortonese, 1940; Eschmeyer et al., 1996).

Alburnus punctulatus Kessler, 1877, described from the Kura River at Tiflis (= Tbilisi) and Borzhom, Georgia, is a synonym. A syntype of Alburnus punctulatus from the St. Petersburg Museum, 84.6 mm standard length, from "R. Kura, Tiflis" is in the Natural History Museum, London (BM(NH) 1897.7.5:34).

Alburnus Brandtii is apparently a manuscript name for this species first reported without a formal description in Brandt (1880) and listed as "Alburnus Brandtii n. sp. 1 ex. Tschaldyr" and attributed to K. Kessler in the account of the travels of Professor A. F. Brandt in Transcaucasia (see Kavraiskii, 1897). Bogutskaya (1997b) lists it as a nomen nudum.

Alburnus microlepis of Kamensky (1901), which is Acanthalburnus microlepis, should not be confused with Alburnus microlepis Heckel, 1843, a distinct species described from Aleppo (= Haleb, Syria).

Key characters

This species is distinguished from A. urmianus by having more lateral line scales, more anal fin branched rays, fewer gill rakers and gill raker morphology according to Saadati (1977). Gill raker counts are the same but scale and anal fin ray counts are generally higher with some overlap. Gill raker morphology does not appear to differ in the fish examined by me. Distribution is the easiest separating factor. Both species are distinguished from other cyprinids in Iran by the dorsal fin spine, 2 rows of pharyngeal teeth, and fin ray and scale counts.

Morphology

Dorsal fin with 3 unbranched and 7-9, usually 8, branched rays, anal fin with 2-4, usually 3, unbranched and 12-19, usually 15-17, branched rays. Pectoral fin branched rays 12-17 and pelvic fin rays 7-9. Lateral line scales 60-87. There is a large pelvic axillary scale. Scales at the base of the anal fin are somewhat enlarged and may be vertically elongate, forming a sheath. The scale focus is sub-central anterior with fine but not numerous circuli and very few posterior radii (less than 10 main radii in the largest fish seen).

Gill rakers 6-12 and sickle-shaped (Saadati, 1977) but this count presumably includes only lower arch rakers. Total gill rakers 10-14, short and only reaching the adjacent raker when appressed. The rounded raker has a triangular flap on its internal surface with the tip of the rounded raker projecting. The raker tip may be squarish or even forked in larger fish. The inner edge of the flap is finely tuberculate. Vertebrae 40-45. Pharyngeal teeth 2,5-5,2 with variants 2,5-5,1, 1,5-5,2, 1,5-5,1, 3,5-5,2, 2,5-4,2, 2,5-4,1, 2,4-5,1, 2,4-4,1 1,5-4,1, 1,5-4,0, 1,4-5,2, 1,4-5,1 and 2,6-5,2. The teeth are hooked at the tip with an elongate flat area below and the largest tooth may be strongly serrated. The posteriormost major row tooth may be almost vertically above the fourth tooth rather than posterior to it. The last unbranched dorsal fin ray is thickened in its lower two-thirds but the last third is thin and flexible. There is an obvious scaleless keel from the pelvic fins to the vent on the belly mid-line. The mouth is oblique and subterminal in adults and most young, oblique and terminal in some young. The gut is relatively short with anterior and posterior loops.

Both males and females, as well as young, may have fine tubercles distributed over the head and especially well-developed ventrally and even on the lips. Belly and lower flank scales have fine tubercles concentrated at the base of the exposed scale, some lining the scale margin. Fine tubercles line the dorsal and ventral surfaces of the pectoral and pelvic fins concentrated on rays but also on membranes, in a single file or variably dispersed.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1) or 8(52); anal fin branched rays 14(4), 15(23), 16(24) or 17(2); pectoral fin branched rays 14(3), 15(36), 16(11) or 17(3); pelvic fin branched rays 8(53); lateral line scales 60(1), 62(2), 63(4), 64(6), 65(11), 66(6), 67(7), 68(4), 69(3), 70(5), 72(1) or 73(2); total gill rakers 10(1), 11(4), 12(18), 13(22) or 14(8); total vertebrae 43(2), 44(10) or 45(13); pharyngeal teeth modally 2,5-5,2(33) with variants 2,5-5,1(6), 1,5-5,2(3) or 1,5-5,1(2).

Sexual dimorphism

Unknown.

Colour

The back and upper head are olive-green to green and the upper flank has a golden sheen. Flanks below are silvery and the abdomen is silvery-white. There is a dark and wide stripe (about orbit diameter) on the flank, not always evident in fresh fish. Above the dark stripe is a narrow golden stripe, about one-third orbit diameter. Dorsal and caudal fins have black tips while paired fins can have a reddish or orange base. The peritoneum is brown with dark blotches or speckles.

Size

Reaches 25 cm.

Distribution

Found in the Kura River of Azerbaijan as far down as Mingechaur but not the lower reaches. In Iran it is found in the Caspian Sea basin including the Aras River shared with Azerbaijan and Iran, as far down as Karadonly, and in the Qarasu a tributary of the Aras. Reported from the Safid River basin (Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Jolodar and Abdoli, 2004; Abdoli and Naderi, 2009) and in the Anzali Talab drainage. Records from the middle Agi Chai or Talkheh River near Tabriz and the Zarrineh River of the Lake Orumiyeh basin are presumably of A. urmianus (Abdoli, 2000).

Zoogeography

The genus and its two species are restricted to the Caspian Sea basin and the adjacent Lake Orumiyeh basin and are presumably derived from a common ancestor related to the Alburnoides-Alburnus lineage.

Habitat

This species inhabits both rivers and lakes.

Age and growth

Females mature at 2 years (Abdurakhmanov, 1962). Spawning probably occurs in the spring judging from fish caught on 31 January which had developing eggs. Türkmen et al. (2001) found fish to 7 years of age in the upper Aras River in Turkey, with three-year-old fish dominant, and also gave length-weight and length-age relationships. Females attained a greater age and size than males.

Food

Food includes aquatic insects, crustaceans and snails, and detritus.

Reproduction

Fecundity is up to 19,060 eggs and egg diameter to 1.87 mm. In Armenia maturity is reached at the end of the second year or beginning of the third year at 80-120 mm and spawning takes place in late April to early May and may continue to late August (Pipoyan and Arakelyan, 1999). In the Turkish Aras, maturity for both sexes began at age 2 years, with all fish mature at 4 years, and spawning started in early May and continued to the end of July. Fecundity reached a mean value of 9705 eggs and egg size reached 1.65 mm (Türkmen et al., 2001).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, few in numbers, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian sea basin.

Further work

The biology of this species has not been investigated and its population biology is also unknown.

Sources

Type material: See above, syntype of Alburnus punctulatus (BM(NH) 1897.7.5:34).

Iranian material: CMNFI 1970-0522, 2, 55.1-71.3 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0536, 4, 70.9-109.3 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0538, 1, 70.7 mm standard length, Gilan, Qezel Owzan River (ca. 36º44'N, ca. 49º24'E); CMNFI 1970-0583, 11, 39.0-79.9 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1979-0454, 8, 37.7-64.7 mm standard length, Zanjan, Qezel Owzan River at Gilavan (36º47'N, 49º08'E); CMNFI 1979-0455, 7, 50.2-123.3 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0695, 15, 71.6-112.7 mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E); CMNFI 1980-0116, 1, 75.5 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 2007-0087, 4, mm standard length, Azarbayjan-e Khavari, Qareh Su (38º22'N, 48º19'E).

Comparative material: CMNFI 1980-0807, 2, 138.2-143.8 mm standard length, Turkey, Ölçek Suyu (no other locality data); CMNFI 1986-0007, 1, 132.2 mm standard length, Turkey, Kars River (ca. 41º00'N, ca. 43º00'E).

Acanthalburnus urmianus
(Günther, 1899)


Gadarchai, October 2011, courtesy of K. Abbasi

 Common names

None.

Systematics

Originally described in the genus Abramis Cuvier, 1816, Berg (1948-1949) placed this species in the genus Alburnoides Jeitteles, 1861 but Saadati (1977) places the species in this genus.

The type material in the Natural History Museum, London comprises 2 specimens, 54.9-58.6 mm standard length, from the Urmi River (BM(NH) 1899.9.30:116-117), 1 specimen, 111.7 mm standard length, from the Ocksa River (BM(NH) 1899.9.30:118) (these three fish being labelled paralectotypes, with 118 being the lectotype, by P. M. Bănărescu in 1980), and 8 specimens, 50.5-111.7 mm standard length, from the Ocksa River (BM(NH) 1899.9.30:119-126), these being syntypes. The series 118-126 were grouped in one jar (seen in May 2014). Günther (1899) refers to the type series as "Five specimens from the Gader Chai and two small ones from the Urmi River; the largest is only 144 millim. long" so there is some confusion over this material.

Key characters

This species is distinguished from A. microlepis by having fewer lateral line scales, fewer anal fin branched rays, more gill rakers and gill raker morphology according to Saadati (1977). Gill raker morphology does not appear to differ in the fish examined by me. Gill raker counts are the same but scale and anal fin ray counts are generally lower with some overlap. Distribution is the easiest separating factor.

Morphology

Dorsal fin with 3 unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 10-13 branched rays. Pectoral fin branched rays 14-16 and pelvic fin branched rays7-8. Lateral line scales 50-68. Scales bear only a few posterior radii and have a subcentral anterior focus. A pelvic axillary scale is present. Gill rakers 10-14, short not quite or just reaching the adjacent raker when appressed; rounded with a projected tip and distinct from its congener according to Saadati (1977) but closely resembling the structure seen in A. microlepis according to my observations (see above under A. microlepis). Pharyngeal teeth usually 2,5-5,2 or 2,5-4,2 with variants 2,4-5,2, 1,5-4,2 or 2,4-4,2. Posterior teeth are hooked at the tip, anterior teeth being rounded, and have no, slight, moderate or even strong serrations. There is a narrow and slightly concave surface below the tip. Some fish have the anterior margin of the concave surface higher than the posterior margin, but this is variable and in some teeth the condition is the reverse. The ventral keel extends from the anus to the base of the pelvic fins and is fleshy from half way to the whole length. The intestine is an elongate s-shaped with a small anterior loop. Total vertebrae 41-43.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1), 8(20) or 9(1); anal fin branched rays 10(1), 11(4), 12(11), or 13(6); pectoral fin branched rays 14(4), 15(17) or 16(1); pelvic fin branched rays 7(3) or 8(19); lateral line scales 50(1), 52(2), 53(2), 55(1), 56(1), 57(2), 59(2), 60(3), 61(3), 62(2), 63(1), 64(1) or 68(1); total gill rakers 10(1), 11(2), 12(6), 13(8), or 14(5); pharyngeal teeth 2,5-5,2(1), 2,5-4,2(1), 2,4-5,2(1) or 2,4-4,2(1); total vertebrae 41(5), 42(7) or 43(2).

Sexual dimorphism

Male fish bear tubercles but fully tuberculate fish have not been examined. One male, 94.7 mm standard length, had a single row of tubercles on anterior pectoral fin rays.

Colour

Overall colour is silvery with a greenish-olive back and flanks with numerous minute brown pigment spots which are crowded above the lateral line to form an inconspicuous darker stripe along the whole side. The dorsal, caudal and pectoral fins have a light to evident speckling of melanophores on the rays and membranes but are almost immaculate in preserved specimens. Larger fish have pigment proximally on the anterior anal fin rays. The peritoneum is silvery but densely speckled with melanophores.

Size

Reaches 15.6 cm standard length, almost 20 cm in total length.

Distribution

This species is endemic to the Lake Orumiyeh basin, apparently in southern and western tributaries (Günther, 1899) although records of A. microlepis from the middle Agi Chai or Talkheh River near Tabriz are presumably of A. urmianus (Abdoli, 2000).

Zoogeography

The closest relative and only congener, Acanthalburnus microlepis, is found in the Caspian Sea basin. Connections between the Lake Orumiyeh basin and the Caspian Sea basin have been suggested by Saadati (1977), an early one in the Pliocene to early Pleistocene resulting in endemic species and a later one in the late Pleistocene resulting in species which are the same as the Caspian or only subspecifically distinct. This species presumably dates from the earlier connection (but see the Lake Orumiyeh drainage basin account for more details).

Habitat

Details of habitat requirements are unknown but is has been collected in both river and lakes.

Age and growth

Fish are mature at 14.4 cm. This species is relatively fast-growing, short-lived species with males attaining 6+ years and females 7+ years in the Kazemi Dam on the Zarrineh River (Abdoli et al., 2008). The von Bertalanffy growth curve was estimated as K = 0.427 in males and 0.506 in females, indicating that females grow faster. The sex ratio was 598♂:912♀ and there were no significant differences between males and females in the linear length-weight relationships.

Food

Diet is generally unknown and guts examined were empty except for a few plant and crustacean remains.

Reproduction

Reproductive data is unknown although this species probably spawns in the spring as do most members of this family.

Parasites and predators

Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. and  Ergasilus sp. on this species.

Economic importance

None.

Conservation

This species is known only from the type series and a few other specimens in museum collections. Its status is unknown.

Further work

Field work should be carried out to determine the habitat requirements, ecology and numbers of this uniquely Iranian fish.

Sources

Type material: See above (BM(NH) 1899.9.30:116-117,  BM(NH) 1899.9.30:118, BM(NH) 1899.9.30:119-126).

Iranian material: CMNFI 1979-0093, 2, 127.5-130.5 mm standard length, Azarbayjan-e Bakhtari, Lake Qowpi (36º57'N, 45º52'E); CMNFI 2007-0098, 1, 156.3 mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0101, 1, 129.3 mm standard length, Azarbayjan-e Bakhtari, Tata'u River south of Miandow Ab (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0105, 1, 90.8 mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); USNM 205904, 1, 84.7 mm standard length, Azarbayjan-e Bakhtari, Nazlu-chay near Rezaiyeh (37º40'N, 45º05'E); USNM 205934, 2, 94.5-141.9 mm standard length, Azarbayjan-e Bakhtari, Lake Qowpi (36º57'N, 45º52'E);  uncatalogued, 4, 105.1-134.9 mm standard length, Azarbayjan-e Bakhtari, Zarineh River (no other locality data);

Genus Acanthobrama
Heckel, 1843

Howes (1981) placed Acanthobrama Heckel, 1843 in the genus Rutilus Rafinesque, 1820 on osteological grounds but most other authors retain Acanthobrama as a distinct genus (Coad, 1984a; Krupp, 1985c; Eschmeyer, 1990; Bănărescu, 1992b) based on the scale, keel and anal fin characters listed below. Durand et al. (2002) include this genus in the Abramis clade based on cytochrome b data. The genus Trachibrama Heckel, 1843 is a lapsus (Krupp and Schneider, 1989).

This genus is characterised by a compressed, deep body of small to moderate size, no barbels, relatively small scales with reduced numbers of radii, a fleshy keel between the base of the pelvic fins and the vent, the last unbranched dorsal fin ray is thickened, spine-like and smooth, and the anal fin is long (9-22 branched rays). Pharyngeal teeth are usually in a single row on each arch. Gut short.

There are 8 species endemic to Southwest Asia with 1 found in southwestern Iran (Goren et al., 1973; Coad et al., 1983; Krupp, 1985c).

Acanthobrama marmid
Heckel, 1843

Gamasiab River, Karkheh River basin, July 2008, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, July 2008, courtesy of K. Abbasi

Common names

كلاش پا (= kalashpa), شبه ساردين (shebeh sardin = pseudo-sardine or resembling sardine), شبه نازي (= shebeh nazy), mahi sim nama (= bream-like fish).

[semnan arrez; samnan areed; arath (Rahemo et al., 2006); marmid, marmid handscherli (= marmid armed with a dagger), marmid abbiad (= white marmid), marmid asphar (= yellow marmid) or marmid mablue (= swallowing or devouring marmid) at Aleppo, arrhada (= dove, lion!) at Mosul (all these latter Arabic names after Heckel (1843b; 1847a), the conflicting names for arrhada included, and are probably antiquated; Tigris bream].

Systematics

Acanthobrama Arrhada Heckel, 1843, Acanthobrama cupida Heckel, 1843, Acanthobrama marmid morpha elata Berg, 1949 and Acanthobrama marmid orontis Berg, 1949 are synonyms.

The type locality for Acanthobrama Marmid is "Gewässern bei Aleppo", for Acanthobrama arrhada "in Mossul", and for Acanthobrama cupida "in Aleppo" according to Heckel (1843b) and "Flusse Kueik bei Aleppo" in Heckel (1847a). The type locality of Acanthobrama marmid morpha elata is Lake Balikli, 12 km from Erzurum, 8 km from the Karasu River, upper Euphrates, in Turkey. The type locality of Acanthobrama marmid orontis is the upper Euphrates region according to Eschmeyer et al. (1996) (but this is an error, see below).

Details on the syntypes of this species and its synonyms arrhada and cupida in the Naturhistorisches Museum Wien are given by Krupp (1985c). Eight syntypes of marmid measuring 41-144 mm standard length are listed from Mosul (in contrast to Heckel's papers where the type locality is Aleppo), the number of fish agreeing with the catalogue in Vienna. These 8 fish are under NMW 55334. Eschmeyer et al. (1996) do not list these fish as types and the card index in Vienna in 1997 concurs. A further 15 fish are listed by Krupp from the Quwayq near Aleppo: 1 fish, 102 mm standard length (NMW 55342 - not in the 1997 card index; but the following NMW fish are listed), 2, 113-139 mm standard length (NMW 55345), 2, 86-121 mm standard length (NMW 55346), 2, 98-126 mm standard length (NMW 55347), 2, 113-132 mm standard length (NMW 55348), 2, 114-138 mm standard length (NMW 79068), and 4 fish in the Senckenberg Museum Frankfurt, 82-112 mm standard length (SMF 543, formerly NMW). Eschmeyer et al. (1996) list NMW 55345-48 (8), NMW 79068 (2), SMF 543 (4) and in the Rijksmuseum van Natuurlijke Historie, Leiden RMNH 2537 (4) and RMNH 2539 (2) (both formerly NMW) as the type series.

Two syntypes of A. arrhada from Mosul, 85-92 mm standard length, are in the Senckenberg Museum Frankfurt (SMF 411, formerly NMW) (F. Krupp, pers. comm., 1985; 85.7-89.0 mm standard length) while 2 others are in the Naturhistorisches Museum Wien, ca. 150 mm standard length (NMW 55335) and 141 mm standard length (NMW 55336) (Krupp, 1985c). However, the Vienna catalogue lists 6 specimens of A. arrhada and in addition to the above material there is also NMW 55334 (8 fish) tagged as syntypes so there is some confusion in what constitutes the type series. Two possible syntypes are in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2538) (Eschmeyer et al., 1996).

Krupp (1985c) records syntypes of A. cupida, 151 mm standard length, (NMW 55340) and 152 mm standard length (NMW 55341). The Vienna catalogue lists 4 A. cupida which agrees with Heckel's description although I observed only NMW 55340 (1 fish), NMW 55341 (1) and also NMW 55342 (1). Eschmeyer et al. (1996) list NMW 55340-43 (1, 1, 1) as syntypes but the numbers indicate 4 fish. The card index in Vienna in 1997 also lists 55505 (5 fish), one of which is designated as the lectotype.

The 2 syntypes of Acanthobrama marmid orontis are in the Zoological Institute, St. Petersburg under ZISP 6720 from "L. Antioch, 1884, Lortet" according to Berg (1949). This subspecies is distinguished only by larger scales from the typical form but the 2 syntypes examined by me had lost their scales and were difficult to count; one seemed to have a count around 64, not as low as 54-55 recorded by Berg (1949). Krupp (1985c) examined type material and new specimens from the Orontes and found them not to differ from A. marmid from the Quwayq and Tigris-Euphrates basins. He accordingly synonymises Acanthobrama marmid orontis with the type subspecies.

Karaman (1972) considered Acanthobrama arrhada to be a subspecies of A. marmid rather than a synonym based on an unusually strongly ossified spiny dorsal fin ray in the former. Since A. marmid was described from Aleppo (= Halab, Syria) and A. arrhada from Mosul, the synonomy of these two taxa may warrant re-examination.

The fish reported from the Tigris River basin of Iran by Nümann (1966) as Xenocypris macrolepidotus was this species (Zoologisches Institut und Zoologisches Museum, Hamburg catalogue number ZMH H2700 examined by me). Saadati (1977) thought it a new species of Acanthobrama but I disagree.

A hybrid with Chalcalburnus (= Alburnus) mossulensis was reported from the Hawr al Hammar in southern Iraq by Krupp et al. (1992).

Key characters

The characters of the genus distinguish this species from all other Iranian cyprinids.

Morphology

Mouth nearly horizontal to oblique, equal or lower jaw slightly behind the upper. The belly has a fleshy keel where the ventral scales do not meet along the mid-line between the pelvic base and the anus. The last unbranched dorsal fin ray is a thickened, stiff and smooth spine, the rigid part varying from 15 to 26% of standard length. The spine may be strong for much of its length and then abruptly become thin and flexible or it may taper gradually to a flexible tip. Some small fish lack an enlarged dorsal fin spine.

Lateral line scales 53-72, scales above the lateral 10-14, scales between the pelvic fin and lateral line 4-7. There is a pelvic axillary scale. Radii are restricted to the posterior field on scales and are few in number. The focus is subcentral anterior to almost central. Dorsal fin with 3 unbranched rays and 7-9 branched rays. Anal fin unbranched rays 3, branched rays 13-22. Pectoral fin branched rays 12-18, pelvic fin branched rays 7-9. Total vertebrae 38-43 (38(3), 39(3), 40(7), 41(5), 42(7) or 43(1) combining Iranian and Iraqi material).. Gill rakers short with a basal swelling, 2-4 on the upper arch, 0-1 at the flexure and 9-12 on the lower arch. Total rakers 12-17. The rakers reach the one below or to its further base end when appressed. Pharyngeal teeth usually 5-5, with the anterior tooth compressed and bluntly pointed, the remainder bevelled with a cutting edge and a hooked tip. The two anterior teeth are more rounded than the others although the second one may have a slight hook and is bevelled. Tigris River basin fish may have 1-2 teeth in a second row. The gut is an elongate s-shape with a large anterior loop in larger fish. The diploid chromosome number is 2n=50, with the karyotype consisting of 8 metacentric, 13 submetacentric and 4 pairs of subtelocentric to acrocentric chromosomes. The karyotype is nearly identical to other Eurasian leuciscine cyprinids (Gaffaroğlu et al., 2006)..

Different body forms occur in slow-flowing and fast-flowing waters. In the former habitat fish have a deep body, often humped behind the head, while in the latter the body is more streamlined (Karaman, 1972). It seems that A. marmid is founded on the humped form and A. arrhada and A. cupida on the streamlined one.

Meristic values for Iranian specimens are:- dorsal fin branched rays 7(1) or 8(8); anal fin branched rays 13(2), 14(2), 15(4) or 17(1); pectoral fin branched rays 13(2), 14(3), 15(3) or 18(1); pelvic fin branched rays 7(1), 8(7) or 9(1); lateral line scales 54(1), 55(2), 56(2), 58(1), 59(1) or 63(1); total gill rakers 12(1), 14(6) or 17(1); pharyngeal teeth 5-4(1) or 5-5(7); and total vertebrae 38(3), 39(1), 41(2), 42(2) or 43(1).

Sexual dimorphism

Fine tubercles are found over the top, sides and bottom of the head in males. Tubercles line the first, unbranched pectoral fin ray irregularly with up to 2 branching rows. Very fine tubercles are found on the adjacent membrane and on the lower pectoral fin surface. Tubercles line the pelvic fin rays in branching rows. The lower caudal fin rays are lined with tubercles. Anterior upper flank scales, all belly scales and lower caudal peduncle scales have their margin lined with tubercles, the peduncle with some tubercles on the mid-scale and the belly with a concentration on the scale base.

Colour

The overall colour is silvery to whitish with the head and back reddish-brown. The flanks can be greyish to blackish from numerous melanophores. There may be a well-developed mid-flank stripe or it may be poorly developed or evident only posteriorly. The pelvic fins are bright red, the pectoral and anal fins less red and the dorsal and caudal fins reddish proximally and black distally. Fin colours may be more orange or yellow than red. All fin rays and membranes have melanophores and these can be quite concentrated such that some fish have dark fins. Young fish in preservative have numerous, distinctive, small to minute, rounded, square or oblong patches of pigment in 1-3, irregular, mid-flank rows. Peritoneum black, silvery with a dorsal concentration of melanophores or with widely scattered melanophores so it appears silvery.

Size

Reaches 20.8 cm (Berg, 1949)

Distribution

This species is found in the Tigris-Euphrates basin of Turkey, Syria, Iraq and Iran, the Quwayq (= Kueik) and Orontes rivers, and possibly the Amik Lake and the Bardan suyu (= stream) near Tarsus (Ladiges, 1960; - Krupp (1985c) suggests these latter should be checked). In Iran it is found in the Tigris River basin including the upper reaches of the Karkheh, the Gav Masiab, the Qara Su, and in marshes such as the Hawr Al Azim.

Zoogeography

The majority of species are found in the Levant which once had connections to the Tigris-Euphrates basin (Krupp, 1985c).

Habitat

Hussain et al. (1997) report this species to be dominant in the small fish assemblages in the Shatt Al-Arab near Basrah, Iraq at 70.8% of 14,084 fish caught. It favours side branches off the Shatt al Arab, presumably to avoid predators which are found in deeper water. Younis et al. (2001b) noted that this species dominated in the polluted and disturbed environment of a dockyard on the Shatt al Arab. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (Hussain et al., 2006) and is also known from large rivers and dams. Also recorded from the Hawr al Azim marsh in Iran.

Age and growth

Al-Nasiri and Salman (1977) studied this species in the Little Zab River, Iraq. Their largest specimen was 13.7 cm. They described length-weight relationships and condition factors but some important length groups were missing from their samples. Condition factor showed a gradual decrease with increasing length and the means for actual and calculated weights were 1.141 and 1.118 respectively. Relative condition factor was 1.0009. Younis et al. (2001) examined three populations of this species in the Shatt al Arab, Iraq and found the 0+ age group to be represented by fish 2.1-11.0 cm long and 1+ age group by fish 8.3-14.1cm. The length-weight relationship was W = -3.821 L2.32. Four age groups with a length range of 4-19 cm were found in the Qarmat Ali River of southern Iraq, with maturity in the first year (Saud, 1997).

Ünlü et al., (1994) examined a population of this species in the Tigris River, Turkey and gave figures for growth in length and in weight. Females grew faster and are larger in size than males at the same age, particularly for age groups III and IV. Condition factor for males was 1.554 and for females 1.550. They found 5 age groups with age group III dominant for both sexes. Overall sex ratio was 1.83 females:1 male. Sexual maturity was attained by 75% of females and 85% of males in the second year of life and all fish in age group III were mature.

Food

Heckel (1843b) suggests that they are ravenous feeders based on the name "swallowing marmid". Gut contents are crustaceans, insects, and plant and gastropod shell fragments in Iranian specimens. Younis et al. (2001a; 2001b) found Shatt al Arab, Iraq fish to be detritivores, having organic detritus as the dominant gut content, followed by phytoplankton (blue-green algae and diatoms), small crustaceans (ostracods, cyclopoids, cladocerans), and aquatic plants, with dominance varying by month. Gut contents were crustaceans, insects, and plant and gastropod shell fragments in fish from Iran examined by me. In a study of the recovering Hammar Marsh, Iraq, diet was 70.77% insects and 9.81% algae with diatoms, plants, crustaceans and snails at less than 10% each, in the Hawr al Hawizah 66.4% insects and 14.1% algae, with amounts of diatoms and various crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 62.7% insects and 17.7% algae with diatoms, plants and various crustaceans at less than 10% each (Hussain et al., 2006).

Reproduction

Younis et al. (2001) found most females to be ripe in March and July samples, and some were spent. Well-developed testes are noted in fish caught on 16 May in Turkey and 7 July near Ravansar, Kermanshahan indicating either a prolonged breeding season or local variations.

Ünlü et al., (1994) report spawning in May to late June for their Tigris River, Turkey population. They cite data for a Keban Dam population (on the Euphrates River in Turkey) where the spawning season is extended and runs from April to August. Egg diameter exceeds 1.2 mm and egg numbers reach 8125, and elsewhere may reach 11,000 eggs. In the Qarmat Ali River in southern Iraq, fecundity reached 1759-9293 eggs.

Parasites and predators

None reported from Iran.

Economic importance

None in Iran. In the early 1990s in Iraq, this species was used for human consumption and for fish meal (Younis et al., 2001).

Conservation

This species is rarely reported from Iranian waters and its status needs to be assessed through further field work. Endangered in Turkey (Fricke et al., 2007).

Further work

Additional field work is required to secure more materials and assess conservation status and biology.

Sources

Type material: See discussion above. Syntypes of Acanthobrama marmid (NMW 55345, NMW 55346, NMW 55347, NMW 55348, NMW 79068, SMF 543); syntypes of Acanthobrama marmid orontis (ZISP 6720), syntypes of A. arrhada (SMF 411, NMW 55335, NMW 55336, NMW 55334); syntypes of A. cupida (NMW 55340, NMW 55341, NMW 55342, NMW 55505).

Iranian material: CMNFI 1979-0287, 2, 89.9-92.1 mm standard length, Kermanshahan, spring near Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0360, 1, 40.6 mm standard length, Khuzestan, Karkeheh River canal (31º40'N, 48º35'E); CMNFI 1979-0377, 2, 28.5-34.6 mm standard length, Khuzestan, Karkheh River (ca. 32º57'N, ca. 47º50'E); CMNFI 1979-0384, 1, 23.1 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1991-0154, 1, 113.6 mm standard length, Khuzestan, Hawr-al-Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0128, 1, 113.6 mm standard length, Kermanshahan, Sarab-e Sabz `Ali Khan (34º25'N, 46º32'E); CMNFI 2007-0114, Kermanshahan, Qareh Su basin (ca. 34º28'N, ca. 46º54'E); ZMH H2700, 1, 145.0 mm standard length, Kermanshahan, Gharasu-Gamasiab-Seymarreh (Qareh Su, Gav Masiab and Simareh rivers, no other locality data); uncatalogued, 1, 101.7 mm standard length, Kermanshahan, sarabs near Kermanshah (no other locality data).

Comparative material: BM(NH) 1931.12.21:22-25, 4, 65.7-84.6 mm standard length, Iraq, Mosul (ca. 36º20'N, ca. 43º08'E); BM(NH) 1974.2.22:1084-1091, 7, 105.1-118.3 mm standard length, Iraq, Najab Bazar (no other locality data); BM(NH) 1974.2.22:1094, 109.3 mm standard length, Iraq, Great Zab River at Aski Kalak (36º16'N, 43º39'E); BM(NH) 1971.4.2:7, 96.5 mm standard length, Iraq, River Tigris near Mosul (ca. 36º20'N, ca. 43º08'E); BM(NH) 1974.2.22:1078-1083, 6, 105.2-122.8 mm standard length, Iraq, Najab Bazar (no other locality data); BM(NH) 1974.2.22:1092, 109.5 mm standard length, Iraq, Najab Bazar (no other locality data); CMNFI 1987-0017, 3, 83.8-108.3 mm standard length, Iraq, Hawr al Hammar (no other locality data); BM(NH) 1920.3.3:147-156, 15, 29.5-102.0 mm standard length, Syria, Ouadi Khneizer (no other locality data); BM(NH) 1968.12.13:108-112, 1 (of 5), 112.6 mm standard length, Syria, Ouadi Khneizer, Khabour (no other locality data) (collections amalgamated as BM(NH 1968,12.13:105-341 seem to include the preceding and following collectiosn, 224 (7 as alizarin specimens), 24.1-69.7 mm standard length); BM(NH) 1968.12.13:113-118, 6, 56.5-117.4 mm standard length, Syria, River Euphrates at Houreira (no other locality data); ZSM 26136, 5, 55.3-80.3 mm standard length, Syria, Assad Reservoir, Euphrates basin (no other locality data); CMNFI 1980-0810, 2, 114.8-118.3 mm standard length, Turkey, Göksu in Tigris River basin (no other locality data); CMNFI 1980-1036, 1, 101.5 mm standard length, Turkey, Keban Dam on Murat Nehri near Elâzığ (no other locality data).

Genus Alburnoides
Jeitteles, 1861

This genus is found in Europe, Asia Minor and Central Asia with ?11 species, with 6 reported in Iran.

The riffle minnows are similar in appearance to the genus Alburnus but have smooth rather than serrated pharyngeal teeth. Arguably this distinction is insufficient to warrant a separate genus but it is retained here as this has not been investigated in depth and the genus has widespread usage. Certainly it is not uncommon to find individuals of Alburnus hohenackeri lacking serrations on their pharyngeal teeth.

Pharyngeal teeth in Alburnoides are in 2 rows with strongly hooked tips but unserrated, scales of medium size, no groove before the dorsal fin, a keel behind the pelvic fins is usually scaleless but may be wholly scaled, short dorsal and moderate to long anal fin, last dorsal fin unbranched ray thickened, decurved lateral line often with a characteristic spotting pattern above and below each pore, and gill rakers short and few.

Alburnoides bipunctatus (Bloch, 1782) was the name applied to most populations across Europe and the Middle East from France north of the Alps eastwards to the Black, Caspian and Aral Sea basins but ongoing research is revealing a greater diversity (Bogutskaya and Coad, 2009; Coad and Bogutskaya, 2009; Seifali et al., 2012).

A record of A. bipunctatus from a qanat at Hormak (29°58'N, 60°51'E) in the Sistan basin by Saadati (1977) is probably an error of labelling or sorting. It is not mentioned in the collector's (R. J. Behnke) original field notes nor in a typed version. Also this species was not collected there by me.  

Records of parasites for fish identified as "A. bipunctatus" in Iran are as follows:

Jalali and Molnár (1990a) record the monogeneans Dactylogyrus alatus and D. chalcalburni from this species in the Zayandeh Rud. Gussev et al. (1993b) also reports the latter species and locality. The monogenean Diplozoon paradoxum is recorded from this species in the Tajan River, Mazandaran (Iranian Fisheries Research and Training Organization Newsletter, 6:7, 1994). Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus ellipsoides. Masoumian et al. (2005) report the protozoan parasites Ichthyophthirius multifilis, Trichodina perforata and Chilodonella, sp. from this species in water bodies in West Azarbayjan. Mortazavi Tabrizi et al. (2005) record Ligula intestinalis in this species from the Sattarkhan Dam in East Azerbaijan. Pazooki et al. (2005) record Trichodina  perforata from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus vistulae, Gyrodactylus sp. and Paradiplozoon sp. from this fish in Zanjan Province. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus alatus, D. chalcalburni and D. pulcher in the Zayandeh River.

To be assigned: CMNFI 1970-0522, 22, 40.4-80.3 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0536, 3, 71.9-89.6 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0546, 3, 57.1-69.4 mm standard length, Gilan, Safid River canal (no other locality data); CMNFI 1970-0551, 1, 108.4 mm standard length, Gilan, Ghaleh River near Fowman (37º13'N, 49º19'E); CMNFI 1970-0583, 16, 40.7-87.3 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1971-0327A, 6, 59.3-81.0 mm standard length, Gilan Shafa River (37º35'N, 49º09'E); CMNFI 1979-0239, 2, 57.1-79.3 mm standard length, Markazi, Nam River near Firuzkuh (35º43'N, 52º40'E); CMNFI 1979-0439A, 4, 53.4-72.2 mm standard length, Gilan, Shafa River (37º35'30"N, 49º05'30"E); CMNFI 1979-0440, 11, 53.7-88.6 mm standard length, Gilan, Lomir River (37º37'N, 49º02'30"E); CMNFI 1979-0441, 4, 52.4-55.7 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0445, 1, 70.6 mm standard length, Gilan, stream 10 km south of Astara (38º21'N, 48º51'E); CMNFI 1979-0453, 2, 45.8-65.1 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0454, 6, 39.6-56.0 mm standard length, Zanjan, Qezel Owzan River at Gilavan (36º47'N, 49º08'E); CMNFI 1979-0483, 2, 93.0-98.6 mm standard length, Mazandaran, Chashmeh River (37º23'30"N, 55º51'30"E); CMNFI 1979-0493, 11, 51.1-82.8 mm standard length, Mazandaran, Tajan River drainage (36º19'N, 53º23'E); CMNFI 1979-0695, 74, 34.1-71.1 mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 489º24'E); CMNFI 1980-0116, 19, 41.1-70.3 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E).

check all species correctly transposed from Bogutskaya and Coad?

add holotype meristics to paratypes frequency

As A. bip from Atrak, Gorgan Gharasu, Tajan, babol, Jaraz, sardab, Aras, Tonekabon, Pol-e Rud, Safis and Anzali Talab (Abdoli and Naderi, 2009).

As A. bip in Tajan River showed negative allometric growth with W = 0.014L2.901 (Patimar et al., 2012).

Alburnoides eichwaldii
De Filippii, 1863

Common names

خياطه (= khayataeh) or ماهي خياطه (= mahi khayateh, tailor or tailoress fish, possibly from lateral line pattern like stitches), لپك (= lapak in Mazandaran), پرك (= parak in Gilaki), sima, kuli.

[gijovcu in Azerbaijan; vostochnaya bystryanka or oriental bystranka, zakavkazskaya bystryanka or Transcaucasian bystranka, Armyanskaya bystryanka or Armenian bystryanka for A. b. armeniensis, all in Russian; spirlin, riffle minnow or riffle bleak in general].

Systematics

Cyprinus bipunctatus was originally described from the Weser River in Germany. Alburnus Eichwaldii De Filippi, 1863, described from the "Kur presso Tiflis" (= Kura River near Tbilisi, Georgia), was regarded as a Caspian Sea basin subspecies of Alburnoides bipunctatus but Bănărescu (1991) briefly stated that it cannot be distinguished from Alburnoides bipunctatus fasciatus (Nordmann, 1840) of the Black Sea basin. Holčík and Jedlička (1994) considered that the observed variation is clinal and subspecies are not warranted. Reshetnikov et al. (1997) also consider subspecies as disputable. There is another nominal subspecies in the Aras River drainage of Armenia, Alburnoides bipunctatus armeniensis Dadikyan, 1972, from the rivers Arpa, Vorotan, Vedi, Marmarik, Kasakh and their tributaries, now regarded as a synonym of eichwaldii (Bogutskaya and Coad, 2009).

Bogutskaya and Coad (2009) resurrect A. eichwaldii, and it is present in at least in the western part of the Caspian Sea basin, west of the Safid River.

A syntype of Cyprinus bipunctatus described from the Weser River, Germany is in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 3357) (Eschmeyer et al., 1996).

Two syntypes of Alburnus eichwaldi from "Tiflis" are in the Naturhistorisches Museum Wien under NMW 55516 and 4 syntypes are in the Istituto e Museo di Zoologia della R. Università di Torino under MZUT N.677 (Tortonese, 1940; Eschmeyer et al., 1996).

Syntypes of Alburnoides bipunctatus armeniensis are in the Zoological Institute, St. Petersburg under ZISP 37502.

Key characters

See B and C 2009 here and below

The pigmentation along the lateral line is distinctive. Total gill raker counts (5-12) are much less than in Alburnus hohenackeri (16-29, usually 20 or more) which has similar general scale and fin rays counts.

A. eichwaldii differs from the related A. gmelini by having fewer branched anal-fin rays (11-14, modally 12-13, vs. 13-16, modally 14-15; means, 12.2 and 14.3, respectively) and a larger number of total vertebrae (mean 41.3 vs. 40.6, statistically different).

Morphology

see tables for counts?

The original diagnosis of A. eichwaldii gave the following characters: the body is deep, its length exceeds the depth in four times; eye large; dorsal-fin rays branched rays 8; branched anal-fin rays 12; scales in the lateral series 50, 11 scales above and 7 scales below lateral line. The original description may be added to by the following combination of characters: the caudal fin lobes are moderately rounded, the fin is not deeply forked; the ventral keel is commonly scaleless but may be variably scaled (up to completely scaled); the head is commonly deep and the snout is slightly to markedly rounded; the upper jaw is slightly protruding over the lower jaw; the tip of the mouth cleft is slightly below the level of the middle of the eye or at about the lower margin of pupil; the number of dorsal-fin rays is 8, rarely 7 or 9; the number of branched anal-fin rays is (10)11-14 with the modal range of 12-13; pharyngeal teeth are commonly 2.5-4.2 and other variants with four teeth in the longer row of the right ceratobranchial, also, less frequently, 2.5-5.2 or 2.5-5.1; the number of total lateral line scales 44-56 (Dadikyan, 1972, 1973, gives 39-56, averaging 48.7, in A. bipunctatus armeniensis); gill rakers 6-10; the number of total vertebrae is (38, 39)40-43 with a mode of 41; predorsal vertebrae are (12)13-15 with a mode of 14; the number of abdominal vertebrae is (18)19-22 with a mode of 21, and that of caudal vertebrae is 19-22 with a mode of 21; the caudal region is commonly one vertebra shorter than, equal to the abdominal region or one vertebra longer than the abdominal region, and the difference between the abdominal and caudal numbers varies from +3 to -1 with a mode of 0; and the most common vertebral formulae are 21+21, 21+20 and 20+21.

 

 

Dorsal fin with 2-3 unbranched and 6-10, usually 8, branched rays, anal fin with 2-3 unbranched and 10-18 branched rays, usually 12-13 (but see below for Iran). Lateral line scales 41-58. Gill rakers 5-12, usually 7-10. Vertebrae 37-44. Pharyngeal teeth 2,5-4,2, rarely 2,5-5,2, 2,4-5,2, or 1,5-4,2, with variants being 1,5-4,1, 2,5-4,3, 2,3-4,2, 2,4-4,2, 1,5-4,0, and 1,2,5-4,3. The chromosome number is 2n=50 (Klinkhardt et al., 1995).

Meristics in Iranian specimens from the Caspian Sea: dorsal fin branched rays 7(6), 8(121) or 9(3); anal fin branched rays 11(1), 12(26), 13(61), 14(32), 15(9) or 16(1); pectoral fin branched rays 12(3), 13(23), 14(71), 15(24) or 16(9); pelvic fin branched rays 6(3), 7(116) or 8(11); lateral line scales 43(4), 44(5), 45(25), 46(29), 47(23), 48(13), 49(7), 50(10), 51(6) 52(5), 54(1) or 55(2); total gill rakers 6(7), 7(35), 8(57), 9(30) or 10(1); pharyngeal teeth 2,5-4,2(14), 2,5-5,2(1) or 2,4-5,2(2); and total vertebrae 37(1), 38(1), 39(4), 40(49), 41(32) or 42(2).

The chromosome number is 2n=50 and Nazari et al. (2009) give further details.

 

Sexual dimorphism

Abdurakhmanov (1962) reports pelvic fin length greater in males and snout length greater in females for this species in Azerbaijan.

Colour

?There is a characteristic pigmentation along the lateral line with a small spot above, and another below, the lateral line opening on each scale. This only appears in preserved material as live fish are an overall silvery colour. It can be absent, mostly in lake forms. The flank has a blue-grey stripe wider than the eye diameter. Above the lateral line there may be a series of 5-9 black lines formed of triangular blotches and 3-5 similar lines below the lateral line. The back and head are dark olive, almost black, dark green or dark brown. The flank above the lateral line may have purple iridescent tints. The flanks can be a golden yellow. The belly and lower head are pearly-white. The dorsal and caudal fins have some grey pigment or may be dark grey. The bases of the pectoral, pelvic and anal fins have orange to red pigmentation which is not well developed in young. The extent and intensity of this pigment is variable between fins, although in some fish it is equally developed in all these fins.

Size

Distribution

Found in river drainages of the southwestern Caspian coast from the Samur (according to Berg, 1948-1949) down to rivers of the Lenkoran’. The Aras River basin also harbours this species.    

Zoogeography

?This species shows considerable variation over its range from Europe to southern Iran. Dadikyan (1973) demonstrated variability in this species in a mountainous region of Armenia within the Aras River basin. Up to 10 characters could be used to distinguish populations within the same river but taken at different altitudes. Populations at similar altitudes but in different rivers (and habitat types, e.g. rushing rocky streams compared to a bog) also varied but the characters were not necessarily the same as those distinguishing altitudinal variants within one river. Local conditions, such as temperature and flow regime, may govern the characters at any one site. Gene flow may play a part as fish are carried downstream by heavy rainfall. Populations living within the same river are presumably more closely related than populations in different river systems but may show more differences than populations at similar altitudes but which have had no gene flow for long periods. These factors complicate designation of subspecies in this species and accurate analysis requires large series of specimens.

Habitat

?This species inhabits small streams and is less frequent in the main flow of large rivers. In Iran, it is one of two most abundant species in Caspian rivers along with Capoeta capoeta (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998). It prefers well-oxygenated water, low in pollution, with hard stream beds. In laboratory experiments with European specimens, Bless (1996) found that reproduction requires a stream velocity of 0.4 ms-1 and a gravel substrate with a diameter of 2-15 cm which allows interstitial flow.

Age and growth

In Azerbaijan, maturity is attained at 1-2 years and life span is 3 years (Abdurakhmanov, 1962).

Food

Food is taken from the bottom or from the water surface, the former being mostly insect larvae and the latter terrestrial organisms which fall on the water. Abdoli (2000) lists Simuliidae, Plecoptera, Ephemeroptera, Chironimidae and Trichoptera. Diatoms are also found in gut contents (Abdurakhmanov, 1962).

Reproduction

?Spawning takes place in spring (April-June) at 13-15.6°C and adhesive eggs are laid on sand or gravel in fast-flowing water. Fecundity reaches 6496 eggs and egg diameter 2.16 mm (Abdurakhmanov, 1962). Bless (1996) reports multiple spawning over a period of 15 weeks in laboratory conditions.

Parasites and predators

Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish (as A. bipunctatus).

Economic importance

Unknown. A. bipunctatus is listed as important to North Americans (Robins et al., 1991). Importance is based on its use as bait and in textbooks. It is also a known feeder on the larvae of the malaria-carrying mosquito.

Conservation

Lelek (1987) considers A. bipunctatus to be vulnerable to endangered in Europe through pollution and eutrophication. It is listed as Vulnerable in Turkey (Fricke et al., 2007). Kiabi et al. (1999), examining Iranian material, consider A bipunctatus to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. These assessments may apply to the current taxon.

Further work

Other populations in Iran related to this taxon are under study (2009).

Sources

Iranian material: CMNFI 2007-0090, ?

Comparative material: See Bogutskaya and Coad (2009).

Alburnoides holciki
Coad and Bogutskaya, 2012


Holotype


 

Common names

None.

Systematics

The female holotype is in the  under  The species was named after

Key characters

The species is distinguished by a combination of characters which includes a

Morphology

Sexual dimorphism

Colour

Size

Attains

Distribution

Zoogeography

Habitat

 


 

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

The numbers and wider distribution of this species should be researched as it is known from only two localities.

Further Work

See under Conservation. Biology is unknown.

Sources

Type material: See above.

Iranian material:

Alburnoides idignensis
Bogutskaya and Coad, 2009

 


Description of holotype. A ventral keel between the pelvics and the anal fin is scaleless
along about 1/2 of its length. There is a
pelvic axillary scale and scales extend over
the proximal bases of the anal fin forming
a sheath. The upper body profile is convex,
similar to the lower profile. The caudal
fin lobes are rounded, the fin is shallowly
forked. The snout is markedly rounded,
stout. The mouth is small, between terminal
and subterminal; the tip of the mouth
cleft is on a level of the lower margin of the
pupil.


Dorsal fin rays are 3 unbranched and 8½
branched, anal fin rays are 3 unbranched
and 12½ branched, branched pectoral fin
rays are 14, pelvic fin branched rays are 7.
The anal fin origin is somewhat in front of a
vertical from the posterior end of the dorsal
fin base. Total lateral line scales number 45
and those to posterior margin of hypurals
44, scales around caudal peduncle 15, scales
above lateral line to dorsal fin origin are 9,
scales below lateral line to anal fin origin
are 5, scales below lateral line to pelvic fin
origin are 4, and midline predorsal scales
are 19. Pharyngeal teeth 2.5-4.2. Gill rakers
number 7, they are short and stubby, the
longest touching the adjacent one when appressed.
Total vertebrae are 38, comprising
19 abdominal and 19 caudal vertebrae. Predorsal
vertebrae number 11.

Description of paratypes.
The body is moderately compressed,
relatively thick. The caudal fin lobes are
rounded, the fin is shallowly forked. The
ventral keel between the pelvics and anal
fin is variably scaled: completely scaleless
(4), scaled along about ¼-1/3 of its length
(11), scaled along ½ of its length (7), scaled
along about 2/3 of its length (3) or completely
scaled (4). The anal fin origin is in
front of a vertical from the posterior end
of the dorsal fin base. The snout is moderately
stout, rounded. The mouth is almost
horizontal, its position is between terminal
and subterminal; the tip of the mouth cleft
is between a level of the lower margin of the
pupil and a lower margin of the eye. The
junction of the lower jaw and the quadrate
is on about a vertical through the anterior
margin of the pupil.



Dorsal fin unbranched rays 3, branched
dorsal-fin rays 6½ (1), 7½ (2) and 8½ (10)

(7.7, 0.63). Anal fin unbranched rays 3,
branched anal-fin rays 10½ (1), 11½ (8)
12½ (4) (11.2, 0.60). The dorsal fin outer
margin is truncate to markedly convex
and the anal fin outer margin is clearly
concave. Pectoral fin branched rays 12(2),
13(5), 14(4), 15(2) (13.5, 0.97), pelvic fin
branched rays 6(1), 7(12) (6.9, 0.28).
Pharyngeal tooth counts are 2.5-4.2
(20), 2.4-4.2 (5), 2.5-4.1 (2), 2.5-4.3 (2),
1.5-4.2 (1). The lateral line is complete with
none, 1 or 2 unpored scales at the posterior
end of the lateral series; total lateral line
scales 41(3), 42(2), 43(1), 44(4), 45(1),
46(2) (43.3, 1.80); lateral line scales to the
margin of hypurals 39(1), 40(3), 41(2),
42(1), 43(3), 44(3) (41.9, 1.77). Scales
around caudal peduncle 12(1), 13(-), 14(3),
15(5), 16(-), 17(4) (15.2, 1.52); scales between
dorsal fin origin and lateral line 8(1),
9(11), 10(1) (9.0, 0.41); scales between anal
fin origin and lateral line 4(5), 5(7), 6(1)
(4.7, 0.63); scales between pelvic fin origin
and lateral line 4(10), 5(3) (4.2, 0.44), and
predorsal scales 17(1), 18(2), 19(6), 20(1),
21(2), 22(1) (19.3, 1.38). Total gill rakers in
the outer row on first left arch number 6(2),
7(3), 8(7), 9(1) (7.5, 0.88). Total vertebrae
38(1), 39(11), 40(1) (39.0, 0.41).
Other characters as in holotype.
Paratypes bear pigmentation above and
below the lateral line pores, forming a pale
line margined with dark although this is
obscured by background pigment on the
caudal peduncle. A mid-flank stripe is diffuse
posteriorly and fades anteriorly. A thin
dark stripe at the junction of the hypaxial
and epaxial muscles masses is evident
but also fades anteriorly. The pigment on
scales above and below the lateral line can
be strongly or weakly expressed, forming
stripes, but can be absent. The back is dark
and obscures a predorsal and postdorsal
stripe. A series of strong melanophores is
present on the inner margin of the pectoral
fin unbranched ray. Most fins lack much
pigment, the dorsal fin pigment lining the
rays being the strongest apart from that
noted on the pectoral fin.
Summarized data for the paratypes and
additional material of A. idignensis material
(excluding holotype).
Dorsal fin unbranched rays 3, branched
dorsal-fin rays 6½ (1), 7½ (10), 8½ (50),
9½ (1); among 46 radiographed specimens
6½ (1), 7½ (10), 8½ (35) (7.7, 0.49) (Table
1). Anal fin unbranched rays 3, branched
anal-fin rays 9½ (1), 10½ (2), 11½ (29),
12½ (23), 13½ (6), 14½ (1); among 46
radiographed specimens 9½ (1), 10½ (2),
11½ (23), 12½ (16) (11.3, 0.67). The dorsal
fin outer margin is truncate to markedly
convex and the anal fin outer margin is
slightly concave. Pectoral fin branched rays
12(2), 13(20), 14(23), 15(15), 16(2), pelvic
fin branched rays 6(3), 7(58), 8(1).
Total lateral line scales 41(4), 42(8),
43(2), 44(10), 45(14), 46(10), 47(7), 48(3),
49(2), 50(1), 51(1); lateral line scales to
the margin of hypurals 39(2), 40(7), 41(5),
42(4), 43(13), 44(9), 45(10), 46(7), 47(2),
48(2), 49(1). Scales around caudal peduncle
12(1), 13(-), 14(11), 15(21), 16(14),
17(12), 18(3); scales between dorsal fin
origin and lateral line 8(3), 9(33), 10(21),
11(5); scales between anal fin origin and
lateral line 4(20), 5(32), 6(8), 7(2); scales
between pelvic fin origin and lateral line
3(2), 4(16), 5(29), 6(15), and predorsal
scales 17(2), 18(6), 19(19), 20(12), 21(12),
22(7), 23(3), 24(1). Total gill rakers in the
outer row on first left arch number 6(7),
7(14), 8(32), 9(8), 10(1).
Vertebral counts given below were calculated
in 46 specimens. Total vertebrae
number (37)38-40 with a mode of 39 (39.0,
0.65) (Tables 2 and 4). Predorsal vertebrae
number 11-13(14) (12.2, 0.4) (Tables 2 and
5). Abdominal vertebrae number (18)19-20
(19.5, 0.55) (Tables 3 and 5). Caudal vertebrae
number (18)19-20 (19.5, 0.55) (Tables
3 and 6). The vertebral formulae are 20+19
(16), 19+20 (14), 20+20 (8), 19+19 (6),
19+18 (1), and 18+20 (1). Thus, the mean
difference between abdominal and caudal
counts varies between +3 and -2 averaging
0 (0.0, 0.88) (Tables 3 and 6).


see tables for counts

Common names

شبه زوري (shebeh zury = resembling zury) in Khuzestan for Alburnoides spp..

Systematics

The holotype (CMNFI 2007-0118) is a male, 106.8 mm TL, 89.2 mm SL from Kermanshahan, Bid Sorkh River between Sahneh and Kangavar, Gav Masiab River drainage, ca. 34°23´N, 47°52´E; 1976 and paratypes (CMNFI 2007-0118A) number 13, 33.5-90.0 mm SL, same data as holotype. The species is named for the Tigris River which was called Idigna in Sumerian (Akkadian: Idiklat; biblical: Hiddekel; Arabic: Dijlah; Turkish: Dicle).

Key characters

This species is distinguished by a combination of characters which includes an unbranched pectoral fin ray strongly lined with melanophores on its inner margin; an eye of an average size, the orbit diameter larger than the snout length and markedly smaller than the interorbital width; caudal fin lobes rounded and fin shallowly forked; a variably scaled ventral keel though most commonly scaled along about 1/3-2/3 of its length; a deep head with a markedly rounded,
stout snout; a small mouth which is between terminal and subterminal; a tip of the mouth cleft on a level from the lower margin of the pupil; commonly 8 branched dorsal-fin rays; 10-12(13-14) branched anal-fin rays; 41-49(50-51) total lateral line scales (39-49 scales to posterior margin of hypurals); commonly 2.5-4.2 or 2.4-4.2 pharyngeal teeth; (37)38-40, with a mode of 39, total vertebrae; 11-13(14) predorsal vertebrae, (18)19-20 abdominal vertebrae; (18)19-20 caudal vertebrae; a caudal vertebral region most commonly one vertebra shorter or one vertebra longer than the abdominal region; the most common vertebral formulae are 20+19 and 19+20, and the difference between  the abdominal and caudal counts averaging 0.

Morphology

Sexual dimorphism

The following characters were significantly different between sexes (p<0.05). Greater in females: head width, postorbital distance, pelvic fin origin to anal fin origin distance. Greater in males: head length, pectoral fin length in pectoral fin origin to pelvic fin origin distance, and pelvic fin length in pelvic fin origin to anal fin origin distance.

Colour

The lateral line is delineated by some darker pigment above and below but not as strongly as in the A. petrubanarescui holotype and obscured by background pigmentation on the caudal peduncle. Some pigment on the flank scales above and below the lateral line give the impression of stripes but is not strongly developed.A mid-flank stripe is not developed. A thin dark stripe separates the epaxial and hypaxial muscle masses. The back is dark and obscures a predorsal and postdorsal stripe. The fins are mostly immaculate, with some melanophores lining the rays of the dorsal and pectoral fins in particular. The unbranched pectoral fin ray is strongly lined with melanophores on its inner margin. The peritoneum is silvery with fine melanophores and some spots.

Size

Distribution

This species is known from some upper reaches of tributaries of Karkheh [Qareh Su] River in the Zagros Mountains. The Karkheh drains into the Tigris just below its confluence with the Euphrates.

Zoogeography

Habitat

This species was captured in the Sarab Dowrah River at an altitude of 1370 m, in clear water at 19°C, with pH 6.8, the shore was bushy, some plants were present in the water, and the river had a stony bed. Other
species recorded together with this species were Barbus lacerta, a “Nemacheilus” sp., Alburnus mossulensis, Capoeta aculeata, Cyprinion macrostomum and Garra rufa.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Comparative material: CMNFI 1979-0278, 5, 43.3-52.8 mm SL, Lorestan, Sarab Dowrah River in Kashkan River drainage, 30 km from Khorramabad (33°34´N, 48°01´E); CMNFI 2007-0075, 36, 38.1-72.1 mm SL, Hamadan, Qareh Su River system, Malayer River at bridge 5 km from Malayer (ca. 34°17´N, 48°47´E); CMNFI 2007-0115, 8, 43.3-62.7 mm SL, Kermanshahan, stream in Karkheh River system north of Kermanshah (ca. 34°34´N, 46°47´E).

Alburnoides namaki
Bogutskaya and Coad, 2009

 

see tables for counts

Common names

None.

Systematics

The holotype, CMNFI 1979-0461, is a female, 91.2 mm SL from Hamadan, qanat at Taveh, 35°07´N, 49°02´E. Paratypes are under  CMNFI 1979-0461A, 188, 27.2-96.9 mm SL, same data as the holotype. The species is named for the Namak Lake. Namak means salt in Farsi.

Key characters

 

Morphology

Sexual dimorphism

The following characters were significantly different between sexes (p<0.05). Head depth, body depth, head width, orbit diameter, and predorsal length were greater in females while pectoral fin length, pelvic fin length, longest dorsal fin ray length, pectoral fin length in pectoral fin origin to pelvic fin origin distance and pelvic fin length in pelvic fin origin to anal fin origin distance were greater in males. One male bore tubercles lining scale margins and sparsely on the top and sides of the head. Tubercles are strongest on scales of the caudal peduncle. The anal-fin rays bear tubercles which follow the branching of the distal rays. Tubercles are present on the dorsal, pectoral and pelvic fin rays but are less developed than those on the anal fin.

Colour

harmonise within?

The lateral line is somewhat darker than the surrounding flank but there are no strong spots or dark outline to canal. Some pigment on flank scales above and below the lateral line give a faint impression of stripes. A mid-flank stripe is only weakly apparent. A predorsal and postdorsal stripe is present on the back. The fins are mostly immaculate, with some melanophores lining the rays of the dorsal and pectoral fins. The flanks were a golden-yellow, belly white, back dark green, base of paired and anal fins orange, other fins hyaline in life. Some paratypes bear strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark. A broad mid-flank stripe can be well developed or weakly expressed and, on the caudal peduncle, obscures the lateral line pigment pattern. However, the lateral line pattern can be weak and this can be seen over the anal fin where the flank stripe does not extend down to the decurved lateral line. The pigment on scales above and below the lateral line (and below the mid-flank stripe) can be strongly or weakly expressed, and in the former case it appears as a series of thin, discontinuous stripes. Some fish have a series of strong melanophores on the inner margin of the pectoral fin unbranched ray. Dorsal fin membranes may be dusky and lack pigment lining the rays. The peritoneum is silvery with a few melanophores.

 

Size

 

Distribution

    

    

Zoogeography

 

Habitat

Habitat data is based only on the collection data. Altitude was 1640 m, water temperature 15.5 °C, pH 6.0, conductivity 1.2 mS, qanat stream width 1.5 m, maximum depth 75 cm, vegetation in water encrusting, shore grassy, gravel and mud bottom, medium current, and water clear in parts, others cloudy and polluted, The species was collected with Capoeta buhsei.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Type material: See above.

Comparative material: CMNFI 2007-0121, 3, 28.0-74.8 mm SL, Hamadan, stream in Qareh Chay basin north of Razan, ca. 35°25´N, 49°02´E; CMNFI 2007-0074, 4, 33.1-41.8 mm SL, Markazi, Qareh Chay, 32 km west
of Arak, 34°03´N, 49°21´E;  ZMH 4183 (7, ).?

Alburnoides namaki
Bogutskaya and Coad, 2009

Diagnosis. The species is distinguished by
a combination of characters which includes
the lack of strong spots or dark outline to the
lateral line canal; a small eye, the orbit width
about equal to the snout length but markedly
smaller than the interidth; caudal
fin lobes rounded and fin shallowly forked; a
sharp scaleless ventral keel behind the pelvic
fins along the abdomen to the anus; a deep
head with a stout snout which is markedly
rounded; a tip of the mouth cleft on the level
below the lower margin of the eye; commonly
8½ branched dorsal-fin rays; 10-13½,
commonly 11-12½, branched anal-fin rays;
(43)44-50(52) total lateral line scales (42-51
scales to posterior margin of hypurals); 2.5-
4.2 pharyngeal teeth (or other variants with
four teeth on the right ceratobranchial);
commonly 39-41 total vertebrae; 11-13(14),
commonly12-13, predorsal vertebrae; 19-
20(21) abdominal vertebrae; 19-21 caudal
vertebrae; a caudal vertebral region most
commonly equal to the abdominal region;
and the most common vertebral formulae are
20+20, 20+19 and 19+20.
Description of holotype. A ventral keel
between the pelvics and the anal fin is completely
scaleless. There is a pelvic axillary
scale and scales extend over the proximal
bases of the anal fin forming a sheath. Dorsal
fin rays are 3 unbranched and 8½ branched,
anal fin rays are 3 unbranched and 12½
branched, branched pectoral fin rays are 13,
pelvic fin branched rays are 6. The anal fin
origin is on a vertical from the posterior end
of the dorsal fin base. Total lateral line scales
number 50 and those to posterior margin of
hypurals 51, scales around caudal peduncle
16, scales above lateral line to dorsal fin origin
are 12, scales below lateral line to anal
fin origin are 6, scales below lateral line to
pelvic fin origin are 7, and midline predorsal
scales are 25. Pharyngeal teeth 2.5-4.3. Gill
rakers number 7, they are short and stubby,
the longest touching the adjacent one when
appressed. Total vertebrae are 40, comprising
20 abdominal and 20 caudal vertebrae.
Predorsal vertebrae number 13.
The upper body profile is convex, similar
to the lower profile. The snout is markedly
rounded, stout. The mouth is small, almost
subterminal; the tip of the mouth cleft is
on a level from the lower margin of the eye.


Description of paratypes. The body is
compressed. The ventral keel between the
pelvics and anal fin is completely scaleless,
very sharp and prominent in all specimens.
The anal fin origin is below the posterior
end of the dorsal fin base. The snout is short
and markedly rounded in smaller and larger
individuals. The mouth is almost subterminal,
with the tip of the mouth cleft on a level
of the lower margin of the eye or below. The
junction of the lower jaw and the quadrate
is on about a vertical through the middle of
the eye.



Dorsal fin unbranched rays 3, branched
dorsal-fin rays 7½ (2), 8½ (48), 9½ (8)
(8.1, 0.41). Anal fin unbranched rays 3,
branched anal-fin rays 10½ (5), 11½ (14),
12½ (29), 13½ (9), 14½ (1) (11.8, 0.88)
(see also Tables 1 and 4 for data based on a
set of another 48 specimens which were radiographed).
The dorsal fin outer margin is
truncate to markedly convex and the anal
fin outer margin is slightly concave. Pectoral
fin branched rays 12(6), 13(33), 14(17),
15(2) (13.3, 0.69), pelvic fin branched rays
6(7), 7(51) (6.9, 0.33).
Pharyngeal tooth counts are 2.5-4.2
(20), 2.4-4.2 (5), 2.5-4.1 (2), 2.5-4.3 (2),
1.5-4.2 (1). The lateral line is complete
with none or 1 unpored scales at the posterior
end of the lateral series; total lateral
line scales 43(1), 44(3), 45(3), 46(11), 47(12), 48(16), 49(8), 50(3), 51(-), 52(1)
(47.3, 1.68); lateral line scales to the margin
of hypurals 42(1), 43(4), 44(5), 45(12),
46(15), 47(10), 48(9), 49(1), 50(-), 51(1)
(46.1, 1.70). Scales around caudal peduncle
14(2), 15(9) 16(13), 17(19), 18(14), 19(1)
(16.6, 1.17); scales between dorsal fin origin
and lateral line 9(4), 10(29), 11(24), 12(-),
13(1) (10.4, 0.70); scales between anal fin
origin and lateral line 4(10), 5(37), 6(10),
7(1) (5.0, 0.65); scales between pelvic fin
origin and lateral line 4(3), 5(31), 6(22),
7(2) (5.4, 0.65), and predorsal scales 18(1),
19(4), 20(13), 21(16), 22(13), 23(4), 24(3),
25(4) (21.3, 1.58). Total gill rakers in the
outer row on first left arch number 5(1),
6(14), 7(26), 8(14), 9(3) (7.0, 0.90).
Vertebral counts were calculated for 48
specimens (including holotype). Total vertebrae
number 39-40(41) (39.7, 0.59) (Tables
2 and 4). Predorsal vertebrae number
(11)12-13(14) (12.2, 0.54) (Tables 2 and
5). Abdominal vertebrae number 19-21with
a mode of 20 (19.8, 0.52) (Tables 3 and
5). Caudal vertebrae number 19-21 (19.9,
0.58) (Tables 3 and 6). The vertebral formulae
are 20+20 (in 21 specimens), 19+20
(10), 20+19 (8), 19+21 (3), 20+21 (3), and
21+19 (2). Thus, the mean difference between
abdominal and caudal counts varies
between +3 and -2 with a mode of 0 (-0.1,
0.92) (Tables 3 and 6).
Many
scales are regenerated in various fish from
this collection, perhaps indicating a traumatic
life.


Comparative remarks. Alburnoides namaki
sp. n. differs from all the congeners primarily
by having a combination of a sharp
scaleles keel, a short markedly rounded
snout, an almost subterminal mouth and a
low number of predorsal vertebrae (modally
12). In tree diagrams based on combined
data (Figs 3-6) A. namaki is clustered
together with A. varentsovi sp. n. from the
northern slope of Kopetdag. Alburnoides
namaki which shares with A. varentsovi sp.
n. (and A. idignensis sp. n.) the lowest number
of predorsal vertebrae (modally 12) is
distinguished by a shallowly forked caudal
fin with rounded lobes (vs. clearly forked,
with pointed lobes), a small, almost subterminal
mouth with the tip of the mouth cleft
on a level from the lower margin of the eye
or below (vs. oblique and terminal, the tip
of the mouth cleft on a level from the middle
of the eye or slightly above), a sharp and
commonly completely scaleless ventral keel
(vs. commonly partly scaled).
 

 

Common names

None.

Systematics

Key characters

 

Morphology

Sexual dimorphism

Colour

Size

 

Distribution

    

    

Zoogeography

 

Habitat

 

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

 

Further Work

 

 

Alburnoides nicolausi
Bogutskaya and Coad, 2009

 

?see tables for counts, check agaisnt etxt


Diagnosis. The species is distinguished
by a combination of characters which includes
an eye of an average size, the orbit
diameter larger than the snout length and
smaller than the interorbital width; caudal
fin lobes rounded and fin shallowly forked;
a variably scaled ventral keel though most
commonly scaled only along about 1/3 of
its length or scaleless; a deep head with a
moderately stout snout which is slightly
pointed; a tip of the mouth cleft on the level
about the lower margin of the pupil, commonly
7½ branched dorsal-fin rays; 8-11½
branched anal-fin rays; (43)43-47(48-50)
total lateral line scales (42-48 scales to posterior
margin of hypurals); commonly 2.5-
4.2 or 2.4-4.2 pharyngeal teeth; commonly
39-40 total vertebrae; 12-13 predorsal vertebrae;
19-20(21) abdominal vertebrae;
18-20 caudal vertebrae; a caudal vertebral
region most commonly one vertebra shorter
than the abdominal region; and the most
common vertebral formulae are 20+19,
19+20 and 20+20.

Dorsal fin rays are 3 unbranched and 7½
branched, anal fin rays are 3 unbranched
and 10½ branched, branched pectoral fin
rays are 13, pelvic fin branched rays are 7.
The anal fin origin is slightly behind a vertical
from the posterior end of the dorsal fin
base. Total lateral line scales number 47 and
those to posterior margin of hypurals 45,
scales around caudal peduncle 17, scales
above lateral line to dorsal fin origin are 10,
scales below lateral line to anal fin origin
are 5, scales below lateral line to pelvic fin
origin are 5, and midline predorsal scales
are 19. Pharyngeal teeth 2.5-4.2. Gill rakers
number 8, they are short and stubby, the
longest touching the adjacent one when appressed.
Total vertebrae are 38, comprising
20 abdominal and 18 caudal vertebrae. Predorsal
vertebrae number 12.


Description of paratypes.

 


Dorsal fin unbranched rays 3, branched
dorsal-fin rays 7½ (52) and 8½ (7) (7.1,
0.33). Anal fin unbranched rays 3, branched
anal-fin rays 8½ (2), 9½ (13), 10½ (32),
11½ (12) (10.0, 0.68) (see also Tables 1 and
4 for 42 radiographed specimens). The dorsal
fin outer margin is commonly truncate,
slightly convex or slightly concave, and the
anal fin outer margin is truncate or only
slightly concave. Pectoral fin branched rays
11(1), 12(30), 13(25), 14(3) (12.5, 0.63),
pelvic fin branched rays 6(6), 7(53) (6.9,
0.30).
Pharyngeal tooth counts are 2.5-4.2
(24), 2.4-4.2 (5), 2.4-5.2 (1). The lateral
line is complete with none, 1 or 2 unpored
scales at the posterior end of the lateral series;
total lateral line scales 42(2), 43(11),
44(15), 45(6), 46(13), 47(7), 48(2), 49(2),
50(2) (45.0, 1.82); lateral line scales to the
margin of hypurals 41(3), 42(14), 43(11),
44(11), 45(8), 46(9), 47(-), 48(2), 49(1)
(43.9, 1.83). Scales around caudal peduncle
13(1), 14(9), 15(30) 16(14), 17(5) (15.2,
0.87); scales between dorsal fin origin and
lateral line 8(2), 9(30), 10(25), 11(2) (9.5,
0.62); scales between anal fin origin and lateral
line 3(1), 4(27), 5(28), 6(3) (4.6, 0.62);
scales between pelvic fin origin and lateral
line 4(21), 5(37), 6(1) (4.7, 0.51), and predorsal
scales 18(4), 19(13), 20(18), 21(10),
22(8), 23(4), 24(2) (20.4, 1.49). Total gill
rakers in the outer row on first left arch
number 5(1), 6(1), 7(33), 8(22), 9(2) (7.4,
0.67).
Vertebral counts given below were calculated
in 42 specimens. Total vertebrae
number 38-40 with a mode of 39 (38.9,
0.58) (Tables 2 and 4). Predorsal vertebrae
number 12-13 (12.6, 0.50) (Tables 2 and 5).
Abdominal vertebrae number 19-21with
a mode of 20 (19.8, 0.53) (Tables 3 and 5).
Caudal vertebrae number 18-20 (19.1, 0.68)
(Tables 3 and 6). The vertebral formulae are
20+19 (in 18 specimens), 19+20 (9), 20+18
(6), 20+20 (4), 19+19 (3), 21+18 (1), and
21+19 (1). Thus, the mean difference between
abdominal and caudal counts varies
between +3 and -1 with a mode of 1 (0.6,
1.08) (Tables 3 and 6).
 

 



 

 

Common names

شبه زوري (shebeh zury = resembling zury) in Khuzestan.

Systematics

The holotype (CMNFI 1979-0281) is a female, 75.0 mm SL, Lorestan, stream in Simareh River drainage, 5 km south of Nurabad (34°03´30´´N, 47°58´30´´E) and paratypes (CMNFI 1979-0281A) comprise 164 specimens, 21.3-65.0 mm SL, same data as holotype. The species is named after a Latin male name Nicolaus, a derivative of the Greek Nikolaos (victory of the people), a compound name composed of the elements nikē (victory) and laos (the people); a Russian name Nikolay and an English name Nicholas, the names of, respectively, Nina Bogutskaya’s elder son and Brian Coad’s son, are also derivatives from Nicolaus.

Key characters

This species differs from all the congeners primarily by having a combination of commonly 7 branched dorsal-fin rays, 8-11 branched anal-fin rays, and 38-40, modally 39, total vertebrae.

Morphology

The body is moderately compressed, relatively thick. The upper body profile is convex similar to the lower profile. The snout is only slightly rounded, almost pointed. The mouth is oblique, slightly below than terminal; the tip of the mouth cleft is slightly below a level of the lower margin of the pupil. The junction of the lower jaw and the quadrate is on about a vertical through the middle of the eye. The caudal fin lobes are rounded, the fin is shallowly forked. A ventral keel between the pelvics and the anal fin is not sharp and is variably scaled: completely scaleless (9), scaled along about ¼-1/3 of its length (9), scaled along  half of its length (6), scaled along about two-thirds of its length (4) or completely scaled (2).  There is a pelvic axillary scale and scales extend over the  proximal bases of the anal fin forming a sheath. The anal fin origin is somewhat behind a vertical from the posterior end of the dorsal fin base.
 

Sexual dimorphism

The following characters were larger in females: pectoral fin origin to pelvic fin origin distance, pelvic fin origin to anal fin origin distance, prepelvic fin length, and mouth width while the following were larger in males: caudal peduncle length, pectoral fin length, pelvic fin length, longest dorsal fin ray length, longest anal fin ray length, pectoral fin length in pectoral fin origin to pelvic fin origin distance, and pelvic fin length in pelvic fin origin to anal fin origin distance.

Colour

?harmonise

The lateral line is delineated by some darker pigment above and below but not as strongly as in A. petrubanarescui holotype and obscured by background pigmentation on the caudal peduncle. Some pigment on the flank scales above and below the lateral line is weak or irregular and an impression of stripes is not very evident. The mid-flank stripe is weak and diffuse, fading anteriorly under the dorsal fin. The back is dark but predorsal and postdorsal stripes are evident. The fins are mostly immaculate, with some melanophores lining rays of the dorsal and pectoral fins in particular. The unbranched pectoral fin ray is lined with melanophores on its inner margin, but not as strongly as in some other samples.

Paratypes can bear strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark, or this pattern may be quite faint. The mid-flank stripe is weak or diffuse and fades anteriorly. A thin line of pigment can be evident separating the hypaxial and epaxial muscle masses, fading anteriorly. The pigment on scales above and below the lateral line (and below the mid-flank stripe) can  be obvious and form a series of thin, discontinuous stripes, or it can be absent. Some fish have a series of strong melanophores on the inner margin of the pectoral fin unbranched ray. The peritoneum is silvery with fine melanophores and some spots.

Size

Distribution

The species is known only from its type locality, a stream in the Simareh River drainage at Nurabad. The Simareh (Seymareh) flows into the Karkheh (Qareh Su) River which enters the Hawr al Hawizeh (Hawr al Azim) on the Iran-Iraq border (Tigris River drainage).          

Zoogeography

In tree diagrams (Bogutksaya and Coad, 2009) based on combined data, this species it is clustered together with another Tigris River basin species, A. idignensis. ??

Habitat

Habitat data is based on collection data. Fish were collected at 2000 m altitude, 19°C water temperature, clear water, pH 6.8, forested shore, stony river bed, moderate amounts of aquatic plants, and no other species taken.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Alburnoides petrubanarescui
Bogutskaya and Coad, 2009

 

?see tables for counts, check against etxt

Holotype. CMNFI 1970-0558, female, 109.1 mm TL, 88.8 mm SL; Iran, Azarbaijan-e Bakhtari, Qasemlou Chay, Orumiyeh [Urmia] Lake basin, ca. 37°21´N, 45°09´E; 27 June 1962; coll. V.D. Vladykov. Paratypes. CMNFI 1970-0558A, 51, 28.7-87.3 mm SL, counts and measurements on 29 fish 33.6-87.3 mm SL; same data as holotype.

Diagnosis. The species is distinguished by a combination of characters which includes a small eye; the orbit width about equal to the snout length but markedly smaller than the interorbital width; caudal fin lobes rounded and the fin shallowly forked; a scaled ventral keel behind the pelvic fins along the abdomen to the anus, a deep head with a stout snout which is markedly rounded; a tip of the mouth cleft on the level below the lower margin of the eye; commonly 7½ (less frequently 8½) branched dorsal-fin rays; 8-10½, commonly 9½, branched anal-fin rays; 44-51 total lateral line scales (42-49 scales to posterior margin of hypurals); 2.5- 4.2 pharyngeal teeth (or other variants with four teeth on the right ceratobranchial); commonly 40-41 total vertebrae; 13-14 predorsal vertebrae; 20-22, commonly 21, abdominal vertebrae; 19-20 caudal vertebrae; a caudal vertebral region most commonly
shorter than the abdominal region; and the most common vertebral formulae are 21+19 and 21+20.
Description of holotype. The caudal fin lobes are rounded and the fin is shallowly forked. A ventral keel between the pelvics and the anal fin is smooth and completely scaled. There is a pelvic axillary scale and scales extend over the proximal bases of the anal fin forming a sheath. The upper body profile is convex, similar to the lower profile. The snout is markedly rounded, stout. The mouth is small, subterminal; the tip of the mouth cleft is on a level below the lower margin of the eye. The body depth enters SL 3.3 times, HL enters 4.3, predorsal length 1.8, caudal peduncle depth 7.7, caudal peduncle length 4.1, length of longest dorsal fin ray 5.2, and length of longest anal fin ray to scale sheath 6.8. Orbit diameter enters HL 3.5 times, snout length enters 3.6, and interorbital width 2.6. Pectoral fin length enters pectoral fin origin to pelvic fin origin distance 1.3 times, and pelvic fin length
enters pelvic fin origin to anal fin origin distance 1.2 times. Dorsal fin rays are 3 unbranched and 7½ branched, anal fin rays are 3 unbranched and 9½ branched, branched pectoral fin rays are 13, pelvic fin branched rays are 7. The anal fin origin is on a vertical from the posterior end of the dorsal fin base. Total lateral line scales number 46 and those to posterior margin of hypurals 45, scales around caudal peduncle 15, scales above lateral line to dorsal fin origin are 9, scales below lateral line to anal fin origin are 5, scales below lateral line to pelvic fin origin are 6, and midline predorsal scales are 21. Pharyngeal teeth 2.5-4.2. Gill rakers number 7, they are short
and stubby, the longest touching the adjacent one when appressed. Total vertebrae are 41, comprising 21 abdominal and 20 caudal vertebrae. Predorsal vertebrae number 13.
The peritoneum is silvery with fine melanophores and some large spots. The lateral line is clearly delineated by darker pigment above and below. Some pigment on flank scales above and below the lateral line give the impression of stripes. A mid-flank stripe is evident. The back is dark and obscures a predorsal and postdorsal stripe. The fins are mostly immaculate, with some melanophores lining the rays of the dorsal and pectoral
fins. The unbranched pectoral fin ray is strongly lined with melanophores on its inner margin.

Description of paratypes.
The body is compressed but relatively thick. The ventral keel between the pelvics and anal fin is not sharp and is completely covered by scales in all specimens. The anal fin origin is below the posterior end of the dorsal fin base. The snout is short and markedly rounded in smaller and larger individuals. The mouth is subterminal, with the tip of the mouth cleft on a level below the lower margin of the eye. The junction of the lower jaw and the quadrate is on about a vertical through the anterior eye margin. The following characters were significantly different between sexes (p<0.05). Greater in females: postorbital length, predorsal length, pectoral fin origin to pelvic fin origin distance, pelvic fin origin to anal fin origin distance. Greater in males: HL, pectoral fin length in pectoral fin origin to pelvic fin origin distance, and pelvic fin length in pelvic fin origin to anal fin origin distance. Dorsal fin unbranched rays 3, branched dorsal-fin rays 7½ (19) or 8½ (10) (7.3, 0.48). Anal fin unbranched rays 3, branched anal-fin rays 8-10½ (9.3, 0.64, including holotype) (Tables 1 and 4). The dorsal fin outer margin is truncate to markedly convex and the anal fin outer margin is slightly concave. Pectoral fin branched rays 13(16), 14(12), 15(1) (13.5, 0.57), pelvic fin branched rays
6(5), 7(24) (6.8, 0.38) Pharyngeal tooth counts are 2.5-4.2 (18), 2.4-4.2 (4), 2.5-4.1 (5), 1.4-4.1 (1), 1.5-4.0 (1). The lateral line is complete with none or 1 unpored scales at the posterior end of the lateral series; total lateral
line scales 43(1), 44(3), 45(2), 46(8), 47(5), 48(6), 49(3), 50(1) (46.7, 1.73); lateral line scales to the margin of hypurals 42(1), 43(3), 44(4), 45(5), 46(6), 47(6), 48(3), 49(1) (45.6, 1.76). Scales around caudal peduncle 14(2), 15(7) 16(10), 17(9), 18(-), 19(1) (16.0, 1.09); scales between dorsal fin origin and lateral line 9(7), 10(18), 11(4) (9.9, 0.62); scales between anal fin origin and lateral line 4(5), 5(23), 6(1) (4.9, 0.44); scales between pelvic fin origin and lateral line 3(1), 4(8), 5(20) (4.7, 0.55); predorsal scales 20(1), 21(10), 22(14), 23(3), 24(1) (21.8, 0.83). Total gill rakers in the outer row on first left arch number 6(3), 7(18), 8(8) (7.2, 0.60). Vertebral counts given below include holotype. Total vertebrae number (39)40-41(42) (40.5, 0.63) (Tables 2 and 4). Predorsal vertebrae number 13-14 with a mode of 13 (13.4, 0.50) (Tables 2 and 5). Abdominal vertebrae number 20-22 with a mode of 21 (21.0, 0.41) (Tables 3, 5). Caudal vertebrae number 19-20(21) (19.5, 0.57) (Tables 3 and 6). The vertebral formulae are 21+19 (in 12 specimens), 21+20 (10), 22+19 (3), 20+19 (1), and 21+21 (1). Thus, the caudal vertebral region is shorter than the abdominal region, rarely equal to it (in 3 specimens), the mean difference between abdominal and caudal counts being +1.4 (std 0.77) (Tables 3 and 6). Other characters as in holotype.
Most paratypes bear strong pigmentation above and below the lateral line pores, forming an evident pale line margined with dark. The broad mid-flank stripe is well-developed. The pigment on scales above and below the lateral line (and below the midflank stripe) form a series of thin, discontinuous stripes. Some fish have a series of strong melanophores on the inner margin of the pectoral fin unbranched ray. The lateral line over the pectoral and pelvic fins can be wavy rather than a smooth decurved line.

Comparative remarks. Alburnoides petrubanarescui sp. n. differs from all the congeners primarily by having a combination of a scaled keel, the lowest number of branched anal-fin rays (modal value 9½ vs. 10½ and more), and the highest value of the difference between the abdominal and vertebral counts. A completely scaled keel is a character shared by A. petrubanarescui sp. n., A. oblongus distributed in the lower reaches of the Syr Darya and Alburnoides sp. from Pulvar (Kor River drainage). However, A. petrubanarescui sp. n. is distinguished from the two other species of this group by having fewer branched dorsal-fin rays (commonly 7½ vs. 8½) and fewer branched anal-fin rays (8-10½ vs. 10-12½). Besides, A. petrubanarescui sp. n. is clearly different from A. oblongus by having larger scales (43-50 total lateral line scales vs. 50-56), 2.5-4.2 and 2.4-4.2 pharyngeal teeth (vs.
2.5-5.2 or 1.5-5.1), fewer gill rakers (6-9 vs. 10-13), a truncate or rounded margin of the dorsal fin (vs. concave). A. petrubanarescui sp. n. differs from Alburnoides sp. from Pulvar, besides some other characters, by fewer dorsal-fin branched rays (commonly 7½ vs. 8½), fewer anal-fin branched rays (8-10½, commonly 9½, vs. 10-12½, commonly 11½) and 21+19 or 21+20 vertebrae (vs. 20+20 or 20+21) the difference between abdominal and caudal counts averaging +1.4 (vs. -0.3).
Etymology. The species is named after the late Petru Bǎnǎrescu, a great freshwater ichthyologist who contributed significantly to our knowledge of fishes of Eurasia.

Distribution. This species is described from a river in the Orumiyeh [Urmia] lake basin and we suppose that it may be an endemic species to the Orumiyeh lake basin.

Habitat data for the type locality (June 1962): water 18 °C, fast current in stream, pebbles and sand bottom, shore grassy, much aquatic plant life, caught with dipnet, other species included Alburnus atropatenae, Barbus
lacerta, “Nemacheilus” sp.

Common names

None.

Systematics

Key characters

Morphology

Sexual dimorphism

Colour

Size

Distribution

    

    

Zoogeography

Habitat

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

Unknown.

Conservation

See under A. eichwaldii.

Further Work

This recently described species needs its conservation status and biology investigated.

Sources

Alburnoides qanati
Coad and Bogutskaya, 2009


Holotype

Kor River, courtesy of Jörg Freyhof
Kor River, courtesy of Jörg Freyhof

Common names

None.

Systematics

The female holotype is in the Canadian Museum of Nature, Ottawa, under CMNFI 1977-0509, 81.5 mm TL, 65.0 mm SL, Fars, at source and along stream of a qanat at Naqsh-e Rostam, Pulvar River system (29°59’30”N, 52°54’00’’E). Paratypes are under CMNFI 1977-0510, 168 (not 178 as in type description) specimens, 24.9-72.5 mm SL, same data as holotype. The species was named after the famous qanat system which taps groundwater to support human survival in desert regions and, incidentally, a habitat for fishes.

Key characters

The species is distinguished by a combination of characters which includes a large eye, the orbit width exceeding both the snout length and the interorbital width, a scaled ventral keel behind the pelvic fins along the abdomen to the anus, commonly 43-47 lateral line scales to posterior margin of hypurals, 2.5-4.2 pharyngeal teeth, commonly 8 branched dorsal fin rays, 10-12 branched anal fin rays, 40-41 total vertebrae, an d the caudal vertebral region equal or longer then the abdominal region (vertebral formulae 20+20 or 20+21).

Morphology

The body is markedly compressed. The upper body profile is convex or, in larger specimens, slightly to markedly straightened while the lower profile is considerably convex. The ventral keel between the pelvics and anal fin is not sharp and is completely covered by scales in all specimens but four possessing a short scaleless portion of keel (about half of keel length) just in front of the anus. The dorsal fin outer margin is truncate to slightly rounded and the anal fin outer margin is truncate to slightly concave. The anal fin origin is behind the posterior end of the dorsal fin base. A pelvic axillary scale is present and the anal fin base is proximally overlain by flank scales. The snout is short and slightly pointed. The mouth is terminal to upturned, with the tip of the mouth cleft on a level from slightly above the middle of the eye to the upper margin of the pupil. The mouth cleft is always turned upward, never horizontal, the lower jaw slightly to moderately projecting relative to the upper jaw, and the junction of the lower jaw and the quadrate is on about a vertical through the anterior eye margin. The lateral line is decurved and only the last few scales are elevated and on the mid-caudal peduncle.

Dorsal fin unbranched rays commonly 3, 4 in 3 specimens only, dorsal fin branched rays 7(3) or 8(28), anal fin unbranched rays 3, anal fin branched rays 10(3), 11(22), 12(6), branched pectoral fin rays 13(4), 14(20) or 15(7), pelvic fin branched rays 7(30).  The lateral line is complete with none, 1 or 2 unpored scales at the posterior end of the lateral series. Lateral line scales to posterior margin of hypurals 41(1), 42(1), 43(5), 44(6), 45(3), 46(8), 47(5) 48(1) or 49(1),  scales above lateral line to dorsal fin origin 9(10), 10(18) or 11(3), scales below lateral line to pelvic fin origin 3(4), 4(20) or 5(7), and scales below lateral line to anal fin origin 4(17), 5(13) or 6(1). Total scale radii 8(1), 9(1), 10(4), 11(8), 12(20), 13(17), 14(16) 15(12), 16(7), 17(3) or 18(1) (13.2, 1.91). Scale radii are restricted to the posterior field encroaching laterally, circuli are eccentric and the focus is anteriorly located. Total gill rakers in the outer row on first left arch 6(4), 7(4), 8(21) or 9(1); gill rakers are very short and widely spaced, not touching the adjacent raker when appressed. Total vertebrae including 4 Weberian vertebrae and last complex centrum 40(14) or 41(17), abdominal vertebrae (including intermediate ones; precaudal vertebrae auctorum) 20 (29) or 21(12), predorsal vertebrae (anterior to first dorsal pterygiophore) 13(24) or 14(6), and caudal vertebrae 20(18) or 21(13). The vertebral formula is 20+20(16), 20+21(12) or 21+20(2). Thus, the caudal vertebral region most commonly (in 93% of examined specimens) is equal to or slightly longer then the abdominal region, the mean difference between abdominal and caudal counts being -0.3. Pharyngeal tooth counts are 2.5-4.2 in 10 fish examined with one additional fish being a variant with 2.4-4.0. Teeth are hooked at the tip and not serrated below it. The gut shape is a simple “S” with an occasional specimen showing a slight flexure to the left of the anterior loop. The general topography of cephalic sensory canals and numbers of pores is typical of most Alburnoides, as described by Bogutskaya (1988). The supraorbital canal is not lengthened in its posterior section and has 7-11, commonly 8-10 pores, with 2-4 (3 in 90%) and 5-7 (6 in 73%) canal openings on the nasal and frontal bones, respectively. The infraorbital canal has 10-15 pores (13 in 38%, 12 in 30%) with 4 (93%) or 5 canal openings on the first infraorbital. The preopercular-mandibular canal is complete, with 11-17, modally 13-16, pores (14 in 38%) with (3)4-6 (5 in 77%) and 7-10 (8 in 62%) canal openings on the dentary and preoperculum, respectively. The supratemporal canal is complete, with (4)5-7 (7 in 54%) pores.

Sexual dimorphism

Head length is longer in males than in females. Pectoral fin length and pelvic fin length are also longer in males.

Colour

Pigmentation of the holotype in 5% formalin consisted of a dark lateral line dividing the hypaxial and epaxial muscle masses and a weakly developed stripe of black pigment on mid-flank prominent posteriorly on the caudal peduncle but fading over the pectoral fin and often interrupted anteriorly. The lateral line pores were lined by pigment dorsally and ventrally. A mid-dorsal line was apparent before the dorsal fin, weakly developed behind the fin. The fins were mostly hyaline with some black pigment lining the fin rays of the dorsal and caudal fins, the dorsal rays of the pectoral fins and the anterior rays of the anal fin.

Overall colouration is silvery with the bases of the pectoral, pelvic and anal fins pink in life. An orange line parallels the anal fin base and the lateral line, lying midway between the two. The ventral surface of the head between the dentaries may be yellow-orange and similarly coloured spots may be found on either side of the dorsal mid-line extending along the whole body. Faint yellow spots occur in rows along the flanks also. Pigmentation in preserved fish is as described for the holotype although the lateral stripe is weakly-developed in some specimens, the mid-flank band of spots of black pigment may be variably developed, and the lateral line may be clearly or only faintly edged by pigment. The peritoneum is rarely dark brown but usually is white-grey to light brown with black spots.

Size

Attains 72.5 mm standard length.

Distribution

Known originally from the Pulvar River drainage of the Kor River basin in southern Iran but also recorded from Harat in the Sirjan basin at 30º01.196'N, 54º20.33'E (material from H. R Esmaeili, 2011; cytochrome b data also places these fish with A. qanati, H. R. Esmaeili, 6 October 2011). 

Zoogeography

This is the southernmost Alburnoides species and may have entered the Kor River basin by headwater capture from the Tigris-Euphrates River basin.

Habitat

The qanat stream in the Pulvar River basin at 15.00 hours on 6 October 1976 had clear and colourless water, a temperature of 21°C, pH 6.8, conductivity 0.475 mS, the current was slow to medium, stream width was about 2 m and maximum depth was up to 1 m, the shore was grassy, plant life in the stream consisted of encrusting and submergent types, and the stream bed was gravel and mud. The Harat locality was at an altitude of 1585 m, pH 8.17, dissolved oxygen 7.25 mg/l, conductivity 816 mS and temperature 22.9-23.3°C.

Type locality, mouth of qanat and stream origin, Brian W. Coad
Type locality, mouth of qanat and stream origin, Brian W. Coad

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

The numbers and wider distribution of this species should be researched as it is known from only two localities.

Further Work

See under Conservation. Biology is unknown.

Sources

Type material: See above.

Other material: CMNFI 1979-0060, 4, 21.0-35.4 mm SL, Fars, spring and irrigation channel, 7 km north of Sa’adatabad (30°06’N, 53°12’E).

Alburnoides taeniatus
(Kessler, 1874)

 

Reported from the Tedzhen River basin (Aliev et al., 1988), Karakum Canal, Kopetdag Reservoir and Uzboi lakes (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) in Turkmenistan on the northeastern border of Iran. It may eventually reach the Caspian Sea basin and the Tedzhen (= Hari) River basin of Iran. No Iranian record.

Genus Alburnus
Rafinesque, 1820

The bleaks and shemayas are found in Europe and the northern parts of Southwest Asia with about 38 species (depending on definitions of the taxa). There are 5 species in Iran. Records of Alburnus orontis Sauvage, 1882 from Iran by Armantrout (1969), Banarescu (1977) and Wossughi (1978) are in error (Krupp, 1985c). Chalcalburnus Berg, 1933 is now regarded as a synonym of Alburnus Rafinesque, 1820. There have been numerous variant views of this synonymy. Bogutskaya (1990) considers Chalcalburnus to be distinct but later, Bogutskaya (1997b; Bogutskaya et al., 2000; Bogutskaya and Naseka, 2004), synonymises it with Alburnus. Reshetnikov et al. (1997) retain Chalcalburnus as a distinct genus as does Eschmeyer in "Catalog of Fishes" (downloaded, 10 August 2007). Banister (1980) points out that the distinction of the genus from Alburnus is based on the relative lengths of the ventral keel and the relative thickness of the last unbranched dorsal fin ray, characters which he views with suspicion in the absence of other corroborating evidence.

This genus is characterised by an elongate, compressed, moderately deep body of small to moderate size, a terminal mouth, no barbels, scales of moderate size, pharyngeal teeth in 2 rows (2,5-5,2 or 2,5-4,2) with hooked tips and usually serrations (often absent), short dorsal fin without a thickened ray, a long anal fin, long and relatively numerous gill rakers, a fleshy keel between the base of the pelvic fins and the vent (the naked part usually not reaching as far forward as the pelvic fin bases in species formerly placed in Chalcalburnus), and a light to brown or black peritoneum. Some authors consider the genus Alburnoides to be synonyms of Alburnus (e.g. Saadati (1977)) while others disagree (e.g. Bogutskaya (1990)). These genera are treated separately here to accord with common usage in Southwest Asia, a conservative measure when there are conflicting opinions.

Jalali et al. (2002) and Jalali and Barzegar (2006) record several parasites from an undescribed Chalcalburnus species in Lake Zarivar, namely Ichthyophirius multifilis, two species of Argulus, a Trichodina species, Dactylogyrus alatus, Diplostomum spathaceum, Myxobolus molnari and Ligula intestinalis. Masoumian et al. (2007) record the myxosporean parasite Myxobolus saidovi from Alburnus maculatus (sic) in the Zayandeh River and Mehdipoor et al. (2004) record the monogenean Dactylogyrus alatus from Alburnus maculatus (sic), also in the Zayandeh River..

Alburnus doriae de Filippi, 1865 has a type locality of "dintorni di Schiraz" but fish resembling this species have not been caught there in late twentieth and early twenty-first century collections. Krupp (1985c) refers 5 specimens from the type series of Alburnus doriae to his Alburnus sellal and 2 specimens to Squalius lepidus. The lectotype (MZUT N.720 or MZUT P1110) of Alburnus doriae is stored in the Istituto e Museo di Zoologia della R. Università di Torino (122.0 mm standard length as measured by me) and 5 paralectotypes (MSNG C.E. 9102) of this nominal species are in the Museo Civico di Storia Naturale di Genova (Tortonese, 1934; 1940; 1961), only one of which is A. doriae (109.1 mm standard length as measured by me). Eschmeyer's "Catalog of Fishes" (downloaded 10 August 2007) has 6 specimens in MSNG C.E. 9102, 5 not this species and gives a locality as probably south of Shiraz.

It seems probable that the fish were collected north of Shiraz, presumably in a Tigris River basin stream based on the other species included in the jar (although Alburnus sellal is more likely to be A. mossulensis, q.v.). These materials may, however, have been mixed and the type locality of this nominal species is obscure.

The species of Alburnus in the Zagros Mountains north of Shiraz and west of Esfahan are currently under investigation and final species identities cannot be given at present. Note that materials identified by Coad (1982d: Alburnus maculatus; 1985: A. doriae) as Leuciscus lepidus were in error. An illustration of A. doriae is given above based on the type material.

Small fishes and members of the genus Alburnus are called kuli in Farsi. In Gilan, kuli are eaten with their heads on and are said to full of phosphorus, conferring open-mindedness, intelligence and sophistication on the Gilanis.

Alburnus atropatenae
Berg, 1925

Ghalechai, Lake Orumiyeh basin, October 2011, courtesy of K. Abbasi
Ghalechai, Lake Orumiyeh basin, October 2011, courtesy of K. Abbasi

Common names

None.

Systematics

The type series is the material called Alburnus filippii by Günther (1899) from "Sujbulak and Superghan near the mouth of the Nazlu Chai" as noted in Berg (1925). This material is in the Natural History Museum, London under BM(NH) 1899.9.30:127, syntype, 1 specimen, 89.7 mm Sl, Azarbayjan-e Bakhtari, Superghan near the mouth of the Nazlu Chai (Sopurghan on the Nazlu Chay is at 37°45'N, 45°12'E); BM(NH) 1899.9.30:128-30, syntypes, 3, 70.7-96.3 mm Sl, Azarbayjan-e Bakhtari, Tatawa Chai near Sujbulak (the Tata'u Chay or Simineh River is not close to Saujbulagh or Mahabad at 36°45'N, 45°43'E so the exact locality of this collection is unclear).

These syntypes bear an external label, apparently in A. Günther's handwriting, listing these fish under the name "brevianalis" which is crossed out and filippii substituted. It appears that Günther originally intended to describe them as distinct and subsequently changed his mind.

Berg's (1925) material was not found in a search of the collections of the Zoological Institute, St. Petersburg (ZISP) in November 1993. Eschmeyer et al. (1996) give the following data: Syntypes: (46) ZIL (ZIL being the old acronym for ZISP) but this material is presumably comparative specimens mentioned by Berg (1925).

Coad and Holčík (1999) demonstrated variation between three populations isolated by the salt Lake Orumiyeh but considered this variation as insufficiently different to warrant taxonomic distinction. Nonetheless, the analysis demonstrated that the three populations have diverged in a measurable manner, presumably through geographical isolation, although ecological factors may have played a part as one sample was from a lacustrine rather than a riverine environment.

Key characters

This species is distinguished from its relatives in the former genus Chalcalburnus (having a short, naked ventral keel) by a combination of characters:-

Species

Total gill rakers

Branched anal fin rays

Pored scales in lateral line

Peritoneum colour

atropatenae

11-16

9-12

46-63

black

chalcoides

18-31

12-19

54-74

light brown

mossulensis

11-18

10-14

58-89

brown to black

tarichi (Lake Van, Turkey)

26-29

9-11

65-82

light brown

Morphology

Dorsal fin rays branched 7-9, modally 8, after 3 unbranched rays, anal fin branched rays 9-12 after 3 unbranched rays, pectoral fin branched rays 13-16 and pelvic fin branched rays 7-8. Lateral line scales 46-63. There is a pelvic axillary scale. The scale focus is slightly anterior or central and there are relatively few anterior and posterior radii about equal in number. The exposed fleshy keel in front of the anus is about 1-4 scales lengths, usually 2, long. Gill rakers lanceolate but short, less than half eye width, reaching between the first and second adjacent rakers or touching the second when appressed, total numbering 11-16. Pharyngeal teeth are hooked at the tip and usually bear a few, large serrations on the larger major row teeth or more rarely have no serrations, apparently size independent. The posteriormost major row tooth may be dorsal rather than posterior to the tooth ahead of it. Tooth counts are usually 2,5-4,2. The gut is an elongate s-shape, sometimes with an anterior loop to the left. Total vertebrae 41-43. Chromosome number is 2n=50 (Nazari et al., 2011).

Meristic values for Iranian material: dorsal fin branched rays 7(2), 8(102) or 9(1); anal fin branched rays 9(5), 10(49), 11(45) or 12(6); pectoral fin branched rays 13(7), 14(44), 15(41) or 16(13); pelvic fin branched rays 7(17) or 8(88); lateral line scales 46(4), 47(5), 48(12), 49(15), 50(13), 51(15), 52(14), 53(5), 54(5), 55(5), 56(2), 58(6) or 63(1); total gill rakers 11(12), 12(30), 13(35), 14(16), 15(7) or 16(2); pharyngeal tooth counts 2,5-4,2(54), 2,4-4,2(2), 2,4-5,2(1), 2,5-5,2(1), 1,5-4,2(1) or 2,5-3,2(1); and total vertebrae 41(4), 42(12) or 43(3).

Sexual dimorphism

Male specimens have small scattered tubercles on the top of the head with fewer tubercles on the side of the head. Tubercles are variably distributed on the head depending on the specimen, or even be different on each side of a single fish. A distinct row may parallel the upper lip, another row may follow the upper eye margin, a patch may be present between the nostril and the upper lip, and there may be tubercles between the mouth and the eye. Very small tubercles line the scale margins on the back, flank and belly and belly scales have a fine row of tubercles on the scale base. Tubercles line the rays of the pectoral, dorsal, pelvic and anal fins and weakly on the caudal fin, the rows branching with the fin rays.

Colour

The back is a dark olive brown to grey, with a narrow stripe. The flank has a dark stripe, as wide as the pupil of the eye, extending onto the head as far as the eye and back to the middle of the caudal fin. The stripe is black to dark green. The flank above the stripe is often lighter in contrast to the darker back and accentuates the distinctiveness of the stripe. The flank below this stripe, the belly and the lower head are silvery, and the stripe is clearly set off from the lower flank. The front of the lower jaw is dark and some of this pigment extends into the floor of the mouth. The iris is silvery on the lower half and dark above. The dorsal fin is faintly pigmented grey along its rays, the caudal fin is grey and the other fins are colourless. Melanophores are present on the dorsal and caudal fin rays and the anterior rays of the pectoral, pelvic and anal fin rays. The nostrils may be dark. The peritoneum is black.

Size

Reaches 21.8 cm.

Distribution

This species is endemic to the Lake Orumiyeh basin and is recorded from the Kazim-chai, Ozband River, Talkheh, Ghalechai, Zarrineh and Tatavi rivers (Günther, 1899; Berg, 1925; Abdoli, 2000; K. Abbasi, pers comm., 2012).

Zoogeography

Lake Orumiyeh was formed during the late Pliocene-Pleistocene, lies at 1275-1295 m, and may well have had a Pleistocene connection to the Caspian Sea basin although this is in dispute (Scharlu, 1968; Schweizer, 1975). Pleistocene shorelines from 30 to 115 m above the present level have been confirmed, and the lake covered twice its present area, but this would not permit an external discharge. Berg (1940) reports benches at levels of about 1800 m, 1650-1550 m and 1500-1360 m, which may represent shorelines, and a level of about 1570 m would have had an outlet to the Aras River basin through the Kara-tepe Pass in the northwest and across the plain near the city of Khvoy. Saadati (1977) suggests two connections with the Caspian Sea, an early one in the Pliocene to early Pleistocene resulting in endemic species and a later one in the late Pleistocene resulting in species which are the same as the Caspian or only subspecifically distinct. A. atropatenae may have its origin in the earlier transgression.

Habitat

Unknown.

Age and growth

Unknown.

Food

Gut contents are insects, crustaceans and worms. Filamentous algae are also present, possibly as accidental inclusions.

Reproduction

Fish captured 25-27 June carried mature eggs.

Parasites and predators

None reported from Iran.

Economic importance

Unknown.

Conservation

Biology is poorly known and numbers and habitat requirements would have to be examined for a conservation assessment.

Further work

The biology of this species requires a detailed study.

Sources

Type material. See above, Alburnus atropatenae (BM(NH) 1899.9.30:127, 1899.9.30:128-30).

Iranian material: CMNFI 1970-0557, 26, 17.9-31.6 mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (ca. 37º27'N, ca. 44º55'E); CMNFI 1970-0558, 8, 25.0-88.7 mm standard length, Azarbayjan- e Bakhtari, Qasemlu Chay (ca. 37º21'N, ca. 45º09'E); CMNFI 1970-0559, 48, 31.4-85.2 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay (37º25'N, 45º10'E); CMNFI 1979-0785, 11, 72.6-123.8 mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (ca. 37º27'N, ca. 44º55'E); CMNFI 1979-0786, 26, 65.0-92.2 mm standard length, Azarbayjan-e Khavari, Guru Lake (37º55'N, 46º24'E); CMNFI 2007-0096, 1, 54.7 mm standard length, Azarbayjan-e Bakhtari, Qasemlu River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); CMNFI 2007-0097, 2, 42.0-54.9 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay basin (ca. 37º16'N, ca. 45º08'E); CMNFI 2007-0103, 6, 43.3-73.3 mm standard length, Kordestan, Zarrineh River basin (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0105, 6, 67.3-112.1 mm standard length, Kordestan, Zarrineh River basin (ca. 36º06'N, ca. 46º20'E); OSU 8122, 2, 73.1-83.5 mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (ca. 37º27'N, ca. 44º55'E); USNM 205904, 2, 73.0-82.6 mm standard length, Azarbayjan-e Bakhtari, Nazlu Chay (37º40'N, 45º05'E); uncatalogued, 1, 81.6 mm standard length, Azarbayjan-e Bakhtari, Haladj River near Mahabad (ca. 36º45'N, ca. 45º43'E) (Coad and Holčík, 1999).

?check against Iraq book

Alburnus caeruleus
Heckel, 1843

Gamasiab River, Karkheh River basin, July 2008, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, July 2008, courtesy of K. Abbasi

Tigris River, Diyarbakır, Turkey, courtesy of Jörg Freyhof
Tigris River, Diyarbakır, Turkey, courtesy of Jörg Freyhof

Euphrates River basin, Adıyaman, Turkey, courtesy of Jörg Freyhof
Euphrates River basin, Adıyaman, Turkey, courtesy of Jörg Freyhof

Common names

None.

Systematics

The type locality is Aleppo (= Halab), Syria and material is held inthe Naturhistorisches Museum Wien. Syntypes are listed in Eschmeyer et al. (1996) as NMW 16688 (4, 65.7-86.6 mm standard length as measured by me), NMW 55511-13 (2, 64.5-75.4 mm standard length, 2, 61.2-71.1 mm standard length, 2, 74.1-77.4 mm standard length), 57161 (3, 59.6-71.1 mm standard length) and additionally ?RMNH 2656 [ex NMW] (4); SMF 100 [ex NMW] (4, 61.3-75.7 mm standard length. See also below.

Key characters

Distinguished from its relatives by ? fewer scales along the lateral line (45-58 compared to 60-89) and a deeper body (2.9-3.5 in standard length compared to 4.0-5.1).

Morphology

Dorsal fin with 3 unbranched and 8-9 branched rays, usually 8, anal fin with 3 unbranched and 13-18 branched rays, mostly 14-16, pectoral fin rays 12-15 and pelvic fin rays 7-8. Lateral line moderately to strongly decurved, scales 43-58. Scales lack radii on the anterior field. The naked ventral keel is obvious. Pharyngeal teeth hooked at tip and deeply notched or serrated below. Modally 2,5-4,2, with variants 2,5-5,2, and 2,5-4,1.Total gill rakers 10-13, just reaching past adjacent raker when appressed. Total vertebrae 39 (Bogutskaya et al., 2000). The body is relatively deep with a slight nuchal hump, 2.9-3.5 times in standard length. The gut is s-shaped. Chromosome number is 2n=50 (Nazari et al., 2011).

Sexual dimorphism

Males have tubercles on the lower jaw, the sides and dorsal surface of the head and on flank scales. Tubercles are evident on the pectoral fin and appear as traces on the pelvic fins.

Colour

Back blackish, flanks silvery. Horizontal stripe along flank sky-blue, more diffuse in larger fish but very evident in smaller ones. Flanks, even lower flanks, and head heavily speckled. The lateral line may bear pigment spots above and below each pore but the stitched effect is not as marked as in some Alburnoides species. Fins generally yellowish, dorsal, anal and pelvic fins apically black to sky blue. The membranes of the dorsal and anal fins are heavily pigmented while the rays are clearer. This pigmentation is more evident anteriorly on small fish but in both large and small fish fins appear dark, especially when the fins are collapsed. On the anal fin, some fish have dark pigment on all membranes, others, even large fish, have less pigmentation distally on the posterior membranes. In larger fish, the pectoral and pelvic fins have dark membranes, the pigmentation fading on the smaller rays. The pectoral and pelvic fins can be orange. In some specimens the edge of the caudal fin is quite dark. The peritoneum is brown to black.

Size

Attains 86.9 mm standard length.

Distribution

Found in the Tigris-Euphrates and Quwayq River systems. The Orontes (= Asi) River is not a locality (Krupp, 1985c). In Iran, it is recorded by Keyvan Abbasi (Iranian Fisheries Research Organization Newsletter, 57:2, 2009) from the Gamasiab and Doab rivers (34º22'16"N, 47º54'51"E at 1412 m altitude and 34º27'11"N, 47º39'34"E at 1322 m) and, given the fishing effort, were quite rare (0.02% of fishing sites, 8 individuals). It may be more widely distributed than museum and literature records suggest.

Zoogeography

The relationships of this species zoogeographically have not been studied.

Habitat

Khalifa (1989) reported this species as widely distributed in rivers and ponds, and it is also found in streams, dams and reservoirs in Iraq. Epler et al. (2001) found it to be the third most dominant species of fish in the Iraqi lakes Habbaniyah, Tharthar and Razzazah, comprising 8.7% of all fish collected.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Large eggs were visible in fish from Syria caught on 19 May, suggesting spring spawning.

Parasites and predators

Unknown in Iran.

Economic importance

None.

Conservation

This species is poorly known and documented in Iran so its conservation status is unknown.

Further work

The biology, distribution and conservation status of this species needs investigation in Iran.

Sources

Type material:- Syntypes NMW 16688, NMW 55511, NMW 55512, NMW 55513, NMW 57161, SMF 100.

?see Excel file where some fish re-identified

Comparative material:-  BM(NH) 1931.12.21:21, 86.9 mm standard length, Mosul, Mesopotamia but this has 17 rakers. ; BM(NH) 1974.2.22:83, 1, 67.9 mm standard length, Iraq, Sirwan, Tigris River near Faish Khabour (no other locality data); ZMB 3364 (possibly syntypes as marked from Vienna Museum), 4, 55.6-65.8 mm standard length, Syria, Aleppo (= Halab); SMF 28638, 14, 69.1-100.7 mm standard length, Syria, Euphrates River, Deir ez zor (35º31'N, 39º54'E); SMF 28678, 3, 59.0-98.8 mm standard length, Syria, Euphrates River upstream Deir ez zor (35º31'N, 39º57'E); SMF 28698, 4, 84.4-105.6 mm standard length,  Syria, Euphrates River, downstream Baath Lake (35º55.723'N, 39º00.572'E); SMF 28712, 3, 51.8-59.3 mm standard length,  Syria, Euphrates River Raqqa to Halebye-Zalebye (35º36.083'N, 39º00.572'E to 53º50.029'N, 39º20.797'E); BM(NH) 1968.12.13:124-135, 40.1-50.7 mm standard length, Syria, Tigris River at Ain Diwar (?); BM(NH) 1968.12.13:147-154, 4, 38.5-71.1 mm standard length, Syria, Quwayq River at Behourte (?).

Alburnus chalcoides
(Güldenstaedt, 1772)

Tajan River, courtesy of Jörg Freyhof
Tajan River, courtesy of Jörg Freyhof

Shafa River, April 2012, courtesy of K. Abbasi
Shafa River, April 2012, courtesy of K. Abbasi

Shafa River, June 2009, courtesy of K. Abbasi
Shafa River, June 2009, courtesy of K. Abbasi

Common names

شاه كولي (shah kuli or shah kooli in Gilaki; kuli is widely used for any small fish and may derive from kul which can mean any pond or sheet of water) or شاه ماهي  (= shah mahi menaing royal fish or king fish in the sense of the best or most important fish); mahi shah kuli; كاس كولي (= kas-e kuli, meaning cup or bowl fish?); aslak in Mazandaran, siah kole (= presumably siah kuli, black fish), safid kuli (= white fish).

[samayi, schamay or schumai, Lankaran samayisi for A. chalcoides longissimus, Kur samayisi for A. chalcoides, all in Azerbaijan; Iranskaya shemaya or Iranian shemaya, Lenkoranskaya shemaya or Lenkoran shemaya, shemaya or shamaya in Russian; Caspian shemaya; bleak, Danube bleak].

Systematics

Cyprinus chalcoides was originally described from the Terek, Sulak and Cyrus (= Kura) rivers, Russia.

Cyprinus clupeoides Pallas, 1776 from the Caspian Sea, Terek and Kura rivers (also spelt clupoides in error), possibly Leuciscus albuloides Valenciennes, 1844 from "rivières de Perse", Alburnus longissimus Warpakhovskii, 1892 from the Geoktapinka River, Lenkoran District, Azerbaijan and Alburnus latissimus Kamenskii, 1901 from the mouth of the Kura River, Azerbaijan are synonyms. Since Alburnus latissimus occurs with Alburnus chalcoides in the Kura River, its status is necessarily equivocal.

Chalcalburnus chalcoides iranicus Svetovidov, 1945 was described as the subspecies of the Iranian shore of the Caspian Sea basin and Alburnus chalcoides longissimus Warpakhovskii, 1892 as the subspecies of the Lenkoran in Azerbaijan neighbouring Iran. Coad (1996b) examined the types of iranicus and longissimus and found them not to be distinguishable. The latter name has priority but both these nominal subspecies, and latissimus, are most probably not distinct from the type subspecies. They were founded on small samples from relatively homogenous spawning populations. Variation may be clinal or related to local temperature and other environmental variables. A very large series of specimens would be necessary to define this.

The Caspian Sea species may be Alburnus chalcoides chalcoides with a distinct subspecies, Alburnus chalcoides mento (Heckel, 1836), in the Black Sea basin although up to 13 subspecies are named from Anatolia and the basins of the Black, Caspian and Aral seas.

The type material of Chalcalburnus chalcoides iranicus is in the Zoological Institute, St. Petersburg (ZISP 31231, holotype (see below), and 3 paratypes 142.0-199.9 mm standard length), the type locality being "a small stream near the hospital near Shahi, Talar River basin" on labels in the Zoological Institute, St. Petersburg and "a small river in the vicinity of town Shakhi (basin of the river Talar, running into the Caspian Sea west of the Gorgan Bay" (Svetovidov, 1945b). Shahi or Qa'emshahr is at 36°28'N, 52°53'E. Svetovidov (1945b) lists the holotype as a female of total length 263.5 mm and body length 226 mm but the holotype in ZISP is 216.7 mm standard length (Coad, 1996b).

The type material of Alburnus longissimus is in the Zoological Institute, St. Petersburg (ZISP 8653, 2 syntypes, 164.8-185.9 mm standard length, from "Fl. Geoktapinka" (Lenkoran). The locality is probably near Prishib at 39°08'N, 48°36'E (Coad, 1996b). ZISP 8654 (6 fish, 121.2-164.4 mm standard length) from the type locality are listed as types in Berg (1911-1914) but not in the ZISP catalogue. Also an A. longissimus syntype from St. Petersburg is in the Natural History Museum, London from "R. Geotapinka" (BM(NH) 1891.10.7:28).

Bagherian and Rahmani (2007; 2009) examined two populations, from the Haraz River and the Shirud, morphometrically. The males and the females between the two populations were different, but this was attributed to environmental factors. Truss analysis separated the two populations. Rahmani et al. (2007) were able almost to separate the two populations using meristic characters. Rahmani et al. (2006) were able to separate populations from the Gazafrud and Haraz rivers using morphometric characters but not meristic ones. Rahmani et al. (2009) used the 18S rRNA gene and found populations from the Haraz, Shirud and Gazafrud rivers were homogenous.

A hybrid of Alburnus chalcoides and Vimba vimba persa was reported from the Safid River (Petrov, 1926) and a hybrid between Leuciscus (= Squalius) cephalus and Alburnus chalcoides is reported from Turkey (Ünver and Erk'akan, 2005; Ünver et al., 2008).

Key characters

The short, naked ventral keel, usually 8 branched dorsal fin rays, distribution, and the characters in the table under A. atropatenae can be used to identify this species.

Morphology

Lateral line scales 54-74. The dorsal and ventral scale margins are parallel or rounded and the anterior margin is wavy or has a pronounced central protuberance. The posterior scale margin can be rounded and more or less smooth or rounded and finely crenulate. Crenulation may be related to size or sexual maturity but is not always evident even in spawning males. Circuli are numerous and fine, radii are few and present on the anterior and posterior fields (a few fish had some scales with no anterior radii), and the focus is slightly subcentral anterior. There is a well-developed pelvic axillary scale. The ventral keel is only naked near the vent and rarely may be scaled along its entire length although Kottelat and Freyhof (2007) have an exposed keel of 8-12 scale lengths, up to 80% of the anus to pelvic fin base distance. Dorsal fin with 2-3, usually 3, unbranched and 7-9, usually 8, branched rays, anal fin with 3 unbranched and 12-19 branched rays, pectoral fin branched rays 13-16, and pelvic fin branched rays 7-9. Ginzburg (1936b) gives counts of 13(7), 14(34), 15(52), and 16(7) for anal fin rays from Iranian material, modally different from my counts below (possibly the last two rays were counted separately but variation between samples is also possible). Gill rakers 18-31, serrated medially and elongate, reaching the second or third adjacent raker when appressed. Total vertebrae 43-45. Pharyngeal teeth 2,5-5,2, more rarely 2,5-5,1, 2,5-5,3, 2,5-5,4, or 3,5-5,3. Teeth are elongate, slender, curved inward, strongly hooked at the tip and strongly serrated with serrations on the anterior margin of the long, narrow and concave grinding surface. The most posterior main row tooth may lie medial to the second tooth. The swimbladder is pointed posteriorly (rounded in Alburnus hohenackeri and A. filippii). The gut is an elongate s-shape. Total vertebrae 41-45.

Meristics in Iranian specimens: dorsal fin branched rays 7(3), 8(55) or 9(2); anal fin branched rays 12(1), 13(4), 14(33), 15(19) or 16(3); pectoral fin branched rays 13(4), 14(9), 15(34) or 16(13); pelvic fin branched rays 7(2), 8(57) or 9(1); lateral line scales 54(1), 55(2), 56(2), 57(5), 58(8), 59(5), 60(14), 61(7), 62(5), 63(6), 64(2), 65(1), 66(1) or 67(1); total gill rakers 18(1), 19(5), 20(12), 21(15), 22(14), 23(9), 24(3) or 25(1); pharyngeal teeth 2,5-5,2(30), 2,5-4,2(1), 2,4-5,2(1) or 2,5-5,3(1); and total vertebrae 42(2), 43(9), 44(32) or 45(7).

Sexual dimorphism

Abdurakhmanov (1962) reports the eye diameter and anal fin base to be larger in males on average for fish from the Kura River basin in Azerbaijan. Iranian males taken in July have small tubercles scattered on top of the head and fine tubercles lining the anterior flank scales. Females are larger than males (Bagherian and Rahmani, 2007)

Colour

The overall colour is metallic silvery and the back is a contrasting olive-green. The iris is bright silver. There is no dark band along the sides. The dorsal and caudal fins are greyish and the other fins colourless to whitish. The peritoneum is light brown but with numerous melanophores in contrast to the dark peritoneum in A. mossulensis.

Size

Reaches 50.0 cm and 1.5 kg (Machacek (1983-2012), downloaded 27 July 2012). Shemaya on the Kura River of Azerbaijan are larger than those in the south Caspian, up to 36 cm as opposed to 29 cm.

Distribution

Found from central Europe to the basins of the Black, western and southern Caspian and Aral seas. It is recorded from the entire southern coast of the Caspian Sea and its rivers, including the Atrak, Gorgan, Gharasu, Shafarud, Tajan, Babol, Haraz, Sardab, Aras, Valiabad, Tonekabon, Pol-e Rud and Safid rivers, the Anzali Talab, Gorgan Bay, southeast, southwest and south-central Caspian Sea (Derzhavin, 1934; Kozhin, 1957; Svetovidov, 1945b; Holčík and Oláh, 1992; Shamsi et al., 1997; Abbasi et al., 1999); Kiabi et al., 1999; Abdoli, 2000; Bagherian and Rahmani, 2007; 2009; Patimar et al., 2010; Abdoli and Naderi, 2009; Nikoo et al., 2010).

Alburnus chalcoides aralensis Berg, 1926 is reported from the Karakum Canal in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen River and Caspian Sea basins of Iran.

Zoogeography

A widespread species with numerous nominal subspecies which have not all been fully investigated. It presumably originated as part of a Danubian or Sarmatian fauna and the subspecies have become isolated in parts of this former basin.

Habitat

Young are rheophilous (Abdurakhmanov, 1975). A migration to piedmont and montane zones used to occur before dams and weirs obstructed movements. Some populations are landlocked while others are semi-anadromous. Knipovich (1921) reports this species from depths of 23.8-25.6 m in the Iranian Caspian Sea. Kottelat and Freyhof (2007) record a tolerance of 14‰ salinity. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab. Shape differences found by Bagherian and Rahmani (2007) in two Iranian rivers were attributed to the Haraz River having a muddy estuary, a shallow slope to the bottom, high turbidity and low water flow in contrast to the Shirud which was sandy with high water flow and high clarity. The latter population developed a more slender body due to increased resistance to water flow.

Age and growth

Life span is 5 years with a theoretical limit of 6.5 years in Azerbaijan (Abdurakhmanov, 1975) and at least 5 years in Iran (Holčík and Oláh, 1992) and Turkey (Tarkan et al., 2005). Sexual maturity is attained at 3 years of age in Azerbaijan and growth is most rapid at an age of 2 years, decreasing thereafter because of high natural mortality (Abdurakhmanov, 1975). The fishes on the spring spawning run in the Anzali Mordab are 10.5-29.0 cm standard length, average 14.0 cm, and 2-5 years old with most (63%) fish in age group 3. Males are mature at 2-4 years and females at 3-5 years. Growth is high during the first 3 years of life and then declines (Holčík and Oláh, 1992). Karimpour et al. (1993) found the Anzali Mordab population to be smaller than the Kura River population but the mordab fish showed greater growth after maturation. The spawning migration into the mordab begins in March and peaks in May and at the beginning of June. Length range was 10.0-24.0 cm, average 16.2 cm with a mean weight of 64.7 g. Age composition was 2-5 years with 3-year-olds comprising 62.5% of the fish. Females formed 57% of the migrating fish. Rahmani (2008) investigated this species in the Haraz and Shirud rivers and found maximum length and weight in a 5- (sic) year old female at 251 mm and 96 g, the most abundant age groups were 2+ and 3+ years for males and females respectively, males in the Shirud population were heavier and longer on average in younger ages while differences in females were not significant, and females of the Shirud population has isometric growth while Haraz fish had positive allometry. The von Bertalanffy growth parameters were Lt = [405.9 (1-e-0.1(t+1.54))] for males in Haraz and Lt = [442.6 (1-e-0.1(t+1.43))] for females in Haraz, and  Lt = [359.5 (1-e-0.145(t+1.002))] for males in Shirud and Lt =[405.9 (1-e-0.1(t+1.54))] for females in Shirud. Females had a higher L while K values for males were relatively higher in the two rivers. Rahmani et al. (2009) found growth was better in the Shirud compared with other populations because this river had desirable biological parameters for immigration. Patimar et al. (2010) compared fish from the Siah and Gorgan rivers and found a five-year life cycle, with negative allometric growth for Siah males and positive allometric growth for Siah females and for both sexes in the Gorgan, and sex ratios were unbalanced in favour of females in both rivers. The von Bertalanffy growth parameters were Lt = [370.08 (1-e-0.15(t+0.70))] for males in Siah and Lt = [432.52 (1-e-0.11(t+1.21))] for females in Siah, and  Lt = [371.79 (1-e-0.14(t+0.96))] for males in Gorgan and Lt = [436.10 (1-e-0.11(t+1.34))] for females in Gorgan.

Food

Holčík and Oláh (1992) report a feeding migration in July to September in the western basin of the Anzali Mordab. Gut contents include diatoms and algae, dragonfly larvae, and copepods (Abdurakhmanov, 1962). Iranian fish had plant fragments, sand grains, crustaceans, insect remains and chironomid larvae in gut contents.

Reproduction

This species is an intermittent spawner with three batches of eggs, only two of which are laid at an interval of 18-19 days. Fecundity reaches 54,700 eggs in Azerbaijan but this is less than that of diadromous populations. Egg diameter is up to 1.9 mm. Spawning takes place in the second half of July to the end of August at water temperatures of 18-25°C in the Mingechaur Reservoir in Azerbaijan. Eggs are laid on rocky bottoms in 15-20 cm of water after a migration into streams or on rocky grounds of reservoirs (Abdurakhmanov, 1962; 1975; Elanidze, 1983). There is a spawning migration into the Kura River from October to April, peaking in December-January, with spawning taking place in spring in the upper reaches (Berg, 1959). In Lake Tuş, Turkey spawning occurred in May-June, egg numbers reached 20,971 and average egg diameter 1.05 mm (Balık et al., 1996).

Svetovidov (1945b) considers that Iranian populations (his iranicus subspecies) spawn nearly throughout the year since fish having ripe sex products were caught in both July and February and young were found along the Iranian coast throughout the year. Spawning takes place in the sea, in areas such as Gorgan Bay, and in the lower reaches of rivers. Nikoo et al. (2010) measured serum sex steroids during spawning in the Valiabad River and concluded that this fish may be a multiple spawner. Khaval (1998) reports a spawning migration into the Safid River despite construction, sand removal and pollution. Holčík and Oláh (1992) report a migration into the Anzali Mordab for spawning in late February to early April (but see above; possibly a confusion between the migration at an earlier date than the spawning act). Karimpour et al. (1993) give an absolute fecundity of 6630 eggs in the Anzali Mordab population while mean relative fecundity is 140 eggs/g of body weight. Iranian fish have 1.5 mm eggs as early as 13 March (fish standard length 213.2 mm) and 1.7 mm eggs on 4 June (fish length 154.6 mm) while eggs are only 1.3 mm on 15 July (fish length 142.8 mm). Larger fish may mature and spawn earlier than younger fish. Rahmani et al. (2009) found a peak gonadosomatic index for males in May and for females in early June in the Shirud. Average fecundity was about 3900 eggs with diameter reaching 1.17 mm. Patimar et al. (2010) in their study of Siah and Gorgan River fish found spawning between April and July in the Siah and March and June in the Gorgan, peaking in May in both rivers. Absolute fecundity was up to 38,340 eggs, mean 8426 eggs in the Siah  and up to 17,263 eggs, mean 4215 eggs in the Gorgan. Relative fecundity was up to 599 eggs/g, average 212 eggs/g of body weight in the Siah and up to 696 eggs/g, average 112 eggs/g in the Gorgan. Mean egg diameters were 1.40 mm in the Siah and 1.27 mm in the Gorgan. These differences in life history (see also Age and growth above) were attributed to differing habitat characteristics.

Parasites and predators

Molnár and Jalali (1992) report the monogeneans Dactylogyrus minor, D. alatus and D. vistulae from this species in the Ghasemlu River, an inland watershed, with the latter species also in the Safid Rud. They also describe a new species of monogenean, Dactylogyrus holciki, from this species in the Beshar River of the Persian Gulf drainage, possibly confusing this Caspian Sea basin cyprinid with A. mossulensis. Molnár and Jalali (1992) also record the monogenean Dactylogyrus chalcalburni from the Safid and Zayandeh rivers, although this Caspian Sea basin cyprinid does not occur in the latter locality, possibly again confusing the same species as noted above. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus pseudodispar. The helminths Pentagramma symmetrica and Mazocea alaosa are recorded from the guts of Chalcaburnus tarichi (sic, presumably A. chalcoides) from the Anzali wetland (Ataee and Eslami, 1999; www.mondialvet99.com, downloaded 31 May 2000). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the monogenean trematodes Dactylogyrus chalcalburni and Gyrodactylus sp.. Sattari et al. (2004, 2005) surveyed this species in the Anzali wetlands, recording Anisakis sp. Maleki and Malek (2007) examined fish from the Shirud in the Caspian Sea basin and recorded the digeneans Posthodiplostomum cuticola, Diplostomum spathaceum, Clinostomum complanatum and Allocreadium sp.Sattari et al. (2007) record the nematode Anisakis sp., the digenean Diplostomum spathaceum and the monogenean Dactylogyrus extensus in this species in the Anzali wetland of the Caspian shore. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Argulus foliaceus. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus foliaceus on this species.

Economic importance

The shemaya was a valuable edible fish on the Kura River of Azerbaijan with catches as high as 500 centners (1 centner = 100 kg) prior to construction of the Kura dam. The catch for Azerbaijan in 1933 was 1950 centners or 2,029,000 fish. Catches in the Mingechaur Reservoir formed by the dam were 133 centners in 1972 (Abdurakhmanov, 1975). Reputedly delicious eating (Lönnberg, 1900b). They are fished for on the spawning run when fatty. In Iran they are caught by cast nets in the inlets and outlets of the Anzali Mordab in spring on the spawning run and by gill nets in the western basin on the feeding migration. Holčík and Oláh (1992) report a catch of 956 kg in the Anzali Mordab in 1990 but catches in recent years may have been confused with the exotic Hemiculter leucisculus (Holčík and Olah, 1990).

Conservation

Holčík and Oláh (1992) report a decline in the numbers of this species owing to damming of rivers where it used to spawn. Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include commercial fishing, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran. Endangered in Turkey (Fricke et al., 2007). 

This species has been artificially bred without hormones on the Shirrud with a fertilisation rate of 90-98%. Hatching took 6 days and the hatching rate was 57% (I.F.R.O. Newsletter, 36:4, 2003). On the Tajan River, induction of ovulation has been carried out using LRH-Aa with metoclopramide and carp pituitary extract (Yousefian et al., 2008). Fertilisation rate was 83%, hatching rate 90% and survival of larvae 81%. Shirvani and Jamili (2009) found excessive levels of cadmium and lead in this fish from regions of Anzali where oil ship traffic was highest. Daei et al. (2009) reported on the effects of cadmium and lead on the iron solute in blood.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture and as food. Lelek (1987) classifies this species as vulnerable to endangered in Europe.

Further work

The various subspecies should be examined using molecular techniques and numbers of this species in Iranian rivers monitored for conservation management.

Sources

The types of C. chalcoides iranicus are included in the meristic data for Iranian specimens.

Type material: See above, Chalcalburnus chalcoides iranicus (ZISP 31231), Alburnus longissimus (ZISP 8653, ?8645, BM(NH) 1891.10.7:28).

Iranian material: CMNFI 1970-0531, 4, 64.5-74.9 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0553, 2, 101.9-163.1 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1971-0327A, 2, 54.5-116.9 mm standard length, Gilan, Shafa River (37º35'N, 49º09'E); CMNFI 1979-0081, 7, 77.8-106.5 mm standard length, Mazandaran, Caspian Sea 3 km west of Chalus (36º41'N, 51º24'E); CMNFI 1979-0434, 4, 47.3-154.6 mm standard length, Mazandaran, Shir River (36º51'N, 50º49'E); CMNFI 1979-0435, 1, 170.5 mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0437, 2, 164.5-175.6 mm standard length, Gilan, Safid River 2 km west of Astaneh (37º16'30"N, 49º56'E); CMNFI 1979-0438, 12, 114.9-158.9 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0439, 2, 156.6-173.2 mm standard length, Gilan, Anzali Mordab (ca. 37º27'N, ca. 49º25'E); CMNFI 1979-0441, 1, 109.8 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0443, 1, 159.6 mm standard length, Gilan, river 34 km north of Hashtpar (38º06'N, 48º53'E); CMNFI 1979-0445, 1, 114.9 mm standard length, Gilan, stream 10 km south of Astara (38º21'N, 48º51'E); CMNFI 1979-0455, 1, 88.5 mm standard length, Zanjan, Qezel Owzan River at Gilavan (36º47'N, 49º08'E); CMNFI 1979-0474, 1, 141.0 mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0686, 23, 25.5-111.0 mm standard length, Gilan, Safid River (37º24'N, 49º598'E); CMNFI 1979-0788, 48, 35.2-74.7 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0120, 17, 56.4-69.5 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0123, 2, 97.0-106.4 mm standard length, Gilan, Safid River around Dakha (no other locality data); CMNFI 1980-0126, 3, 182.1-213.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1980-0132, 7, 18.7-142.8 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0142, 2, 135.0-187.2 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1980-0908, 3, 45.4-155.2 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

Alburnus filippii
Kessler, 1877

Qarasu, Aras River basin, January 2011, courtesy of K. Abbasi
Qarasu, Aras River basin, January 2011, courtesy of K. Abbasi

Common names

كولي كورا (= kuli-ye Kura), ماهي مرواريد or مرواريد ماهي (= mahi morvarid or morvarid mahi, meaning pearl fish).

[Kur kumuscasi in Azerbaijan; Kurinskaya ukleika or Kura bleak, ukleika filippi or Filippi's bleak, both in Russian].

Systematics

The lectotype of Alburnus Filippii as designated by N. Bogutskaya is in the Zoological Institute, St. Petersburg (ZISP 2926) and is from "Fl. Kura pr. Tiflis", Acad. Brandt, 1867, 75.3 mm standard length. Paralectotypes are ZISP 2925, 13 fish, same data as lectotype, 43.0-84.4 mm standard length, ZISP 2914, 2 fish, "Fl. Kura pr. Borshoma", Acad. Brandt, 1867, 83.6-87.6 mm standard length, and ZISP 50412, 16 fish, "Reka Kura Tiflis", Acad. Brandt, 1867, 60.6-88.6 mm standard length. A syntype, 57.3 mm standard length, is in the Natural History Museum, London from Tiflis (BM(NH) 1897.7.5:33, formerly in ZISP).

Alburnus filipii var. Kessler in Brandt, 1880 from the Tchaldyr Lake is also this species.

Knipovich (1921) reports a Caspian basin species Alburnus philippii Kessler which is presumably a misspelling of filippii. The specific name is sometimes spelt filippi, which is incorrect.

Abdurakhmanov (1962) compares a sample from the Kura River basin with one from the Kendalanchaya in the Aras River basin of Azerbaijan and finds 15 characters are significantly different on average. Fish from the Kura have a longer head, greater dorsal and anal fin heights, and longer pectoral, pelvic and upper and lower caudal fin lobes while fish from the Aras have more scales in the lateral line, a deeper head, body and caudal peduncle, and a longer anal fin base, pectoral-pelvic fin distance and snout, and a greater interorbital width. No taxonomic status is assigned these two populations.

A hybrid with Alburnus charusini hohenackeri (= Alburnus hohenackeri) was reported by Petrov (1926) from the Safid River and the Kumbashinka in Lenkoran.

Key characters

This species is distinguished from its relative (Alburnus chalcoides, also with a long, naked ventral keel) by having modally 7 branched dorsal fin rays and generally lower anal fin ray counts although these do overlap (10-21, usually 12-16 in Iran for alburnus; 9-13, usually 10-12, for filippii). See also table under A. atropatenae.

Morphology

Dorsal fin with 3 unbranched and 6-8, usually 7, branched rays, anal fin with 2-3, usually 3, unbranched and 9-13 branched rays, usually 10-12. Pectoral fin branched rays 12-16, pelvic fin branched rays 6-8, usually 7. Lateral line scales 46-64. Scales have a wavy anterior margin, an overall vertical oval shape, sometimes tapering to a rounded posterior point and sometimes more rounded, few anterior and posterior radii, and a subcentral anterior focus. There is a pelvic axillary scale. The naked ventral keel usually extends more than half way from the anal papilla to the pelvic fin insertion but is often completely scaled, notably in fish from the Safid River basin. The scaled keel runs from the papilla to the pelvic fin base. Gill rakers 12-21, reaching the second or third adjacent raker when appressed. Pharyngeal teeth 2,5-5,2 (but see below for Iranian specimens) with variants 2,5-5,1, 1,5-5,2, 1,5-5,1, 2,5-4,2, 2,4-5,2, 2,5-4,1, 2,4-4,2, 1,5-4,1, 1,4-5,1, and 1,5-4,2. Teeth are stongly hooked and strongly serrated. Serrations are on the anterior margin of each tooth. The degree of hook and serration development varies individually and does not seem to be size related. Some fish have little development of either character. The area below the hook is an elongate, flat to concave surface. Vertebrae number 38-43. The swimbladder has a rounded end in contrast to the pointed end in Alburnus chalcoides. The gut is an elongate s-shape with a small anterior loop. The chromosome number is 2n=50 and Nazari et al. (2009; 2011) give further details.

Meristic variation in Iranian specimens: dorsal fin branched rays 6(1), 7(44) or 8(5); anal fin branched rays 9(1), 10(19), 11(24), 12(5) or 13(1); pectoral fin branched rays 12(3), 13(19), 14(20), 15(7) or 16(1); pelvic fin branched rays 6(3), 7(42) or 8(5); lateral line scales 46(1), 49(1), 50(5), 51(5), 52(4), 53(12), 54(5), 55(2), 56(5), 57(6), 58(1), 60(2) or 63(1); total gill rakers 12(4), 13(8), 14(19), 15(10), 16(6) or 17(3); pharyngeal teeth 2,5-4,2(10), 2,4-5,2(2), 2,4-4,2(2), 2,5-5,2(1), 1,5-4,2(2), 1,5-5,2(1), 1,5-4,1(1) and 1,4-5,1(1); and total vertebrae 38(2), 39(8), 40(18), 41(9) or 42(1).

Sexual dimorphism

Males and females have moderate-sized tubercles widely scattered on the top of the head, on the snout and lining the lower edge of the jaw. Much smaller tubercles are scattered among the ones on top of the head.

Colour

The back is brown, flanks silvery and the belly white. A characteristic dark streak, as wide as the eye, runs along mid-flank. Fins are hyaline. The peritoneum is brown or light with large scattered melanophores.

Size

Reaches 17.0 cm standard length.

Distribution

Found only in the Caspian Sea basin from the Kura River of Azerbaijan to the Safid River of Iran including headwaters in Turkey, Armenia and Iran at altitudes over 3000 m. It is distributed from the upper to the lower reaches of the Aras (Qareh Su) and Safid (Qezel Owzan) rivers in Iran and in the Anzali Talab (Holčík and Oláh,1992; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli and Naderi, 2009).


Zoogeography

The relationships of this species with other Alburnus needs to be examined. It presumably originated as part of a Sarmatian fauna, isolated in the Caspian Sea.

Habitat

Primarily a freshwater species, this minnow may be found in the brackish outlets of the Anzali Mordab (Holčík and Oláh, 1992). Jolodar and Abdoli (2004) note that it is found more in upstream waters than A. alburnus.

Age and growth

Life span is about 5 years with maturity at 1 year for males and 2 years for females.

Food

Gut contents include plant remains, mayflies and algae (Abdurakhmanov, 1962). Iranian specimens contain insect remains, a few crustaceans and sand grains. One sample from the Qareh Su north of Ardebil had been feeding on water beetles (Hydrophilidae) but also spiders and scarab beetles (Euoniticellus sp.) indicating food is also taken from the surface.

Reproduction

Eggs number up to 14,210 and diameters up to 1.51 mm. May is the principal spawning month in Azerbaijan (Abdurakhmanov, 1962). Male fish caught on 6 June in Iran had tubercles scars on top of the head while female fish from another locality (Zanjan River) taken on 8 June had mature eggs measuring 1.2-1.3 mm. Spawning probably occurs in May and June in Iran, depending on local conditions.

Parasites and predators

Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. elegans in the Beheshtabad River. Mortazavi Tabrizi et al. (2005) record Ligula intestinalis and Bothriocephalus acheilognathi in this species from the Sattarkhan Dam in East Azerbaijan. Undoubtedly food for various predatory fishes. Pazooki et al. (2005) record Trichodina  perforata from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus vistulae and Gyrodactylus sp. from this fish in Zanjan Province.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include medium numbers, habitat destruction, medium range (25-75% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian sea basin. Vulnerable in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs investigation.

Sources

Type material: See above, Alburnus Filippii (ZISP 2926, 2925, 2914, 50412, BM(NH) 1897.7.5:33).

Iranian material: CMNFI 1970-0538, 8, 34.9-61.8 mm standard length, Gilan, Qezel Owzan River (ca. 36º44'N, 49º24'E); CMNFI 1970-0552, 1, 50.1 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1979-0448, 1, 70.9 mm standard length, Azarbayjan-e Khavari, Ahar Chay 8 km from Ardabil (38º18'30"N, 48º22'E); CMNFI 1979-0452, 2, 52.4-54.9 mm standard length, Azarbayjan-e Khavari, Qezel Owxan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 9, 43.7-73.3 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0455, 17, 42.8-62.5 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0695, 3, 61.3-63.5 mm standard length, Gilan, Safid River (36º46'N, 49º24'E); CMNFI 2007-0081, 1, 51.0 mm standard length, Zanjan, Zanjan River near Soltaniyeh (ca. 36º27'N, ca. 48º45'E); CMNFI 2007-0082, 11, 41.2-59.6 mm standard length, Zanjan, Zanjan River basin near Zanjan (ca. 36º36'N, ca. 48º32'E); CMNFI 2007-0087, 6, 55.7-83.1 mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0107, 3, 41.1-42.3 mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E).

Alburnus hohenackeri
Kessler, 1870

Anzali Wetland, November 2011, courtesy of K. Abbasi
Anzali Wetland, November 2011, courtesy of K. Abbasi

Marun River, courtesy of Kai Borkenhagen
Marun River, courtesy of Kai Borkenhagen

Common names

ماهي مرواريد or مرواريد ماهي (= mahi morvarid or morvarid mahi, meaning pearl fish), kuli (= general term for small fish), كولي ايراني (= kuli-ye Irani).

[simali gafgaz kumuscasi for A. c. charusini or zagafgaziya kumuscasi for A. c. hohenackeri, both in Azerbaijan; ukleika or bleak, zakavkazskaya ukleika or Transcaucasian bleak, persidskaya ukleika or Persian bleak, sefidrudskaya ukleika or Safid River bleak, all in Russian; Caucasian bleak (as A. hohenackeri)].

Systematics

Alburnus Hohenackeri was originally decsribed  from Karabakh, Azerbaijan, on the Kura River. The taxon in Iran was formerly included within the wide-ranging species Alburnus alburnus (Linnaesus, 1758). Cyprinus Alburnus was originally described from Europe.

Synonyms are Alburnus charusini Herzenstein in Zograff and Kavraiskii, 1889 described from the Kamysh-Samarskie lakes between the Volga and Ural rivers in Kazakhstan, Alburnus alasanicus Kamenskii, 1901 from the Alasan, Alazan' or Alazani River, a left bank Kura River tributary in Georgia, Alburnus lucidus var. macropterus Kamenskii, 1901 described from the Alazan' River, Alburnus alburnus charusini natio elata Petrov, 1926 from the Prorva River (lower reaches of the Terek River), the Sulak River and the Divichi Liman, western Caspian Sea, Alburnus striatus Petrov, 1926 from "Kizil-Agachskogo Zaliva" (Kizil-Agach Bay, Turkmenistan) and "Astrabadskogo Zaliva" (= Astrabad or Gorgan Bay, Iran), and Alburnus alburnus natio dagestanicus Petrov, 1930 (sic) but later in the same paper given, and probably originally meant, as A. a. charusini n. dagestanicus) described from the "Kaukasusküste des Kaspische Meeres".

Alburnus pseudospirlinus Petrov, 1926 from "Novaya Rechkaya (nizov'ya Sefid-Rud)" (= new stream, lower Safid River) is a hybrid of this species and Alburnoides bipunctatus (sic) (Berg, 1948-1949). A hybrid with Alburnus filippi was described from the Kumbashinka River in the Lenkoran and from the Safid River (Petrov, 1926).

The holotype of Alburnus hohenackeri is in the Zoological Institute, St. Petersburg (ZISP 2839). The holotype of Alburnus charusini is in the Zoological Museum of Moscow State University under MMSU P.1314. Four fish as listed as questionable syntypes under MMSU P.1812 by Svetovidova (1978) although according to Eschmeyer et al. (1996) the original says P.1314 with a unique holotype only.

This species was recognised as Alburnus charusini in Iran but characters overlap with Alburnus alburnus, a highly variable species (Gäsowska, 1974). In any case hohenackeri has priority over charusini. Literature sources conflict on the correct name. Petrov (1926; 1930) refers to Alburnus alburnus hohenackeri Kessler, 1877 for fish in northern Iran with natio persicus Petrov, 1926 in the Safid River, natio dagestanicus Petrov, 1930 in the Dagestan area of Azerbaijan and natio kumbashensis Petrov, 1926 from the Kumbashinka River and Lake Ol'khovskoye in the Lenkoran area of Azerbaijan. Natio are not recognised by the Zoological Code of Nomenclature (Ride et al., 1985). Liška and Pivnička (1985) refer southern and southeastern populations of this species to Alburnus alburnus albidus Costa, 1838, and this would include the Iranian populations. These fish are separated from the type subspecies by having 39-47 lateral line scales, most frequently 42-44 (44-54, most frequently 47-50 in A. a. alburnus), branched anal fin rays 10-17, most frequently 13-15 (14-21, most frequently 16-19), and head length as % of body length 22-27 most frequently 23-25 (19-25, most frequently 21-23). N. Bogutskaya (pers. comm., 1995) and Reshetnikov et al. (1997) refer Iranian fish to Alburnus alburnus hohenackeri as there is a definite character break at the Terek River separating northern populations from southern ones. Petrov (1930) came to a similar conclusion on the name of the Iranian populations in his study as noted above. Aburakhmanov (1962) too refers the taxon hohenackeri to fish found in the Kura and Aras rivers and in rivers of the Lenkoran coast (and presumably the Iranian coast) while his charusini are north of the Apsheron Peninsula. Bogutskaya and Naseka (2004) and Kottelat and Freyhof (2007) recognise A. hohenackeri as a distinct species.

Key characters

This species can be confused with Alburnoides eichwaldii which has similar scale, fin ray and pharyngeal counts. A key distinction is the total gill raker count of 16-29 (usually 20 or more) in this species as opposed to 5-12, usually 7-10 in Alburnoides. Alburnus rakers are more than twice as long as those in Alburnoides and, being more numerous, are crowded on the arch without the large gaps between individual rakers which characterises Alburnoides. Modal dorsal fin branched ray count of 8 separates it from A. filippii. ?separation from other Alburnus in Caspian

Morphology

Dorsal fin branched rays 7-9, usually 8, after 2-4 unbranched rays, anal fin branched rays 10-21 after 3-4 unbranched rays (note that anal fin count will be a narrower range if A. hohenackeri is recognised as distinct from a widespread A. alburnus (see Iranian counts below). Pectoral fin branched rays 11-16 and pelvic fin branched rays 6-9. Lateral line scales 36-55. Scales bear both anterior and posterior radii with a few curved radii in the lateral fields. The focus is subcentral anterior and circuli are numerous and fine. The naked ventral keel is often wholly or partially covered by scales. Gill rakers 15-29, elongate reaching the third, or rarely second, below when appressed. Vertebrae 36-46. Pharyngeal teeth 2,5-5,2 with variants 2,5-5,1, 2,5-5,3, 1,5-5,2, 1,5-5,1, 2,5-4,2, 2,4-5,2, 2,4-5,1, 2,4-4,2, 1,5-4,2, 2,5-4,1, 1,5-4,1, 1,4-4,1. The elongate and narrow teeth bear a strongly hooked tip and have evident serrations in most specimens although some lack them entirely. The gut is an elongate s-shape with a small anterior loop. The posterior end of the swimbladder is rounded (pointed in Alburnus chalcoides). The chromosome number is 2n=50-52, generally 50 (Klinkhardt et al., 1995).

The natio persicus from the Safid River has dorsal fin branched rays 7-9, anal fin branched rays 12-16 and lateral line scales 40-45. Fish from the Kura-Aras basin and Lenkoran (hohenackeri) have anal fin branched rays 10-15, lateral line scales 38-48, pharyngeal teeth 2,5-5,2, total gill rakers 16-25 and total vertebrae 37-42 (courtesy of N. Bogutskaya, Zoological Institute, St. Petersburg).

Meristics for Iranian fish including Petrov's (1930) counts of dorsal and anal branched rays and lateral line scales for Safid River fish are:- branched dorsal fin rays 7(7), 8(76) or 9(8); branched anal fin rays 12(6), 13(37), 14(28), 15(16) or 16(2); branched pectoral fin rays 12(2), 13(18), 14(17) or 15(3); branched pelvic fin rays 7(11) or 8(29); lateral line scales 39(2), 40(8), 41(10), 42(28), 43(13), 44(9), 45(7), 46(1), 47(1), 48(1), or 50(1); total gill rakers 19(1), 20(2), 21(18), 22(7), 23(5), 24(4) or 25(3); pharyngeal teeth 2,5-5,2(13), 2,5-4,2(11), 2,5-4,1(1), 2,4-5,2(2) or 2,4-4,2(1); and total vertebrae 37(2), 38(24), 39(20), 40(7) or 41(1).

Sexual dimorphism

Tubercles line the edge of each scale and in single file line the rays of all fins. Fine tubercles cover the whole head.

Colour

The overall colour is bright silvery with the posterior scale margins grey on the upper flank. The back is dark blue to olive or bluish-green and is sharply distinct from the lighter flanks. The mid-line of the back has a narrow dark line. The lateral line and the area above it have some pigmentation, concentrated along the lateral line itself, but there is no dark stripe or it is only faintly developed and is bluish or greyish. Above this stripe is an iridescent golden-green stripe only visible at a certain angle. The bluish or greyish stripe is more evident in preserved material. The belly and lower head surface are pearly-white. The iris is silvery with a yellow ring along the outer eye rim but very little around the pupil. The upper part of the iris may have some dark pigment. The dorsal and caudal fins have dark rays and transparent membranes but may be a dirty yellow. Membranes may have some pigment, particularly on the dorsal fin. The upper anterior edge of the pectoral fin has a little dark pigment while the rest of the fin is colourless to grey or orange. Some fish have a yellow base to the pectoral fin. The pelvic and anal fins are usually colourless, although the anal rays may have some grey or there may be some yellow, orange or red on the fin generally. The caudal fin tip is dark grey.

In preserved fish, most flank pigment is above the lateral line. Lateral line scales have pigment both above and below the pore so the pore stands out. This is not as distinctive as in some Alburnoides spp.. A mid-dorsal stripe is more evident in smaller fish and is obscured by the generally darker back and upper flank pigmentation in larger fish. The peritoneum is a light silvery with scattered melanophores. A flank stripe may be developed although not as strongly as in Alburnus filippii; the stripe is more a darker area along the muscle mass divide between a lighter upper flank and lower flank.

Size

Reaches 20 cm.

Distribution

Found in the Caspian Sea basin. It is reported from the Aras River (including the upper reaches of its tributary, the Qara Su) to the Atrak River along the Caspian coast of Iran including the Anzali Talab and Gorgan Bay, and the Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Tonekabon; Pol-e Rud and Safid rivers (Derzhavin, 1934; Holčík and Oláh, 1992; Kiabi et al., 1999; Abbasi et al., 1999; Abdoli and Naderi, 2009). Also widely introduced across western, central and eastern Iran, including in the Ab-e Sirvan in the upper Diyala River, Lake Zarivar, in the Zayandeh River of the Esfahan basin, in the Kalshur, Jajarm and Qareh Su of northeastern Dasht-e Kavir basin, and in the Hamun Kushk, and Kahak and Sistan dams of the Sistan basin, and possibly in Minab (= Esteghlal) Dam (A. Abdoli, pers. comm., 1995; J. Holčík, in litt., 1996;.Abdoli, 2000; Ghorbani Chafi, 2000; A. Afzali, pers. comm., 2002; Esmaeili et al., 2011).

Zoogeography

This is a widespread species showing great morphological variability over its range, sometimes recognised as taxa. Zoogeographical relationships of these taxa and of the species to other Alburnus have still to be worked out.

Habitat

This species is found in open waters of lakes along the shore or in slow rivers, avoiding turbid conditions and heavy vegetation. There was a mass mortality, presumed to be of this species, on the Babol Sar beach on 24 June 1963 (USNM 270909). It is found more abundantly at river estuaries along the Iranian Caspian shore than Alburnus filippii (Jolodar and Abdoli, 2004).

Age and growth

Maturity is attained at 3 years and life span is up to 9 years. In more northern waters, most spawning males are 3+ and 4+ years while females are 5+ and 6+ years. Iranian populations probably have a similar structure but the age groups would be lower. Mature males averaged 9.7 cm and females 10.5 cm in one study in Russia (Berg, 1948-1949).

Food

Food is planktonic crustaceans, benthic crustaceans such as amphipods, flying insects which land on the water surface, aquatic insects such as backswimmers (Notonectidae), algae, diatoms, and fish eggs and fry. It is an important prey item for other fishes.

Reproduction

Spawning in Europe takes place from April to July in shallow water over a hard bottom. June is the main spawning month in Azerbaijan judging by egg diameters and condition factors (Abdurakhmanov, 1962). Older fish spawn first. Water temperature is usually at 15-16°C or more. Spawning takes place in 3-6 stages at intervals of 9-11 days. The eggs adhere to stones, branches or vegetation. Fecundity is up to 10,000 eggs and egg diameter to 1.4 mm. Incubation lasts about 1 week. Iranian specimens had 1.1 mm diameter eggs in a sample caught on 11 June and mature males were collected on 10 July. Specimens collected in September showed egg resorption while those taken in December had small, developing eggs and those taken in April with better developed eggs. The specimens were small and spawning probably occurs in July for these fish and possibly June for larger ones.

Parasites and predators

Molnár and Jalali (1992) record the monogeneans Dactylogyrus parvus, D. alatus and D. chalcalburni from Alburnus charusini on the Safid Rud.

Gussev et al. (1993b) report the monogenean, Dactylogyrus chalcalburni, from this species in the Zayandeh Rud but this fish does not occur there. The parasite may have been found in Alburnus mossulensis. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. Barzegar et al. (2008) record eye parasites from this fish including the digeneans Diplostomum spathaceum and Tylodelphys clavata.

Some European populations of Sander lucioperca feed almost exclusively on this species. Spent adults are known to eat their own eggs.

Economic importance

The scales contain silvery crystals of guanine which are extracted and used to make essence d'orient (or pearl essence) for artificial pearls. About 5000 fish are required for 100 g of essence. Schools in the lower Don River of the Black Sea number up to 10 million fish weighing 30 tonnes. This abundant species is of indirect commercial importance as food for more valued fishes but it has also been used as food for humans.

Conservation

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, habitat destruction, widespread range (75% of water bodies), and present in other water bodies in Iran. Endangered in Turkey (Fricke et al., 2007). 

Further work

The biology of this species needs investigation, especially in relation to habitats and other fish species where it has been introduced by accident.

Sources

Iranian material: CMNFI 1970-0510, 8, 44.5-72.1 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0580, 27, 33.9-56.1 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1970-0589, 21, 22.5-67.9 mm standard length, Gilan, Safid River (37º12'N, 49º54'E); CMNFI 1971-0343, 1, 63.5 mm standard length, Gilan, Langarud at Chamkhaleh (37º13'N, 50º16'E); CMNFI 1979-0265, 30, 61.6-90.4 mm standard length, Gilan, head of Anzali Mordab at Abkenar (37º28'N, 49º20'E); CMNFI 1979-0432, 22, 34.4-54.3 mm standard length, Mazandaran, Sardab River branch (36º41'N, 51º22'E); CMNFI 1979-0435, 1, 51.9 mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0480, 6, 14.4-64.3 mm standard length, Mazandaran, Gorgan Rver at Gonbad-e Kavus (37º15'30"N, 55º09'E); CMNFI 1980-0122, 41, 29.8-59.0 mm standard length, Mazandaran, Nerissi River (36º38'N, 52º16'E); CMNFI 1980-0147, 5, 44.3-61.5 mm standard length, Gilan, Lashtenesha River (37º21'N, 49º52'E).

Alburnus mossulensis
Heckel, 1843

Kor River, courtesy of Jörg Freyhof
Kor River, courtesy of Jörg Freyhof

amasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi

Common names

شاه كولي (shah kuli = king fish), shah kuli-ye jonubi (= southern king fish), شاه ماهي (= shah mahi, meaning king fish), shah kuli mosulenzis.

[simnan , semnan or samnan, semnan tuyel; sink, or zurri at Mosul (zurri also used for Chondrostoma regium according to Heckel (1847a), but is also used for Aphanius spp., Gambusia and any small fishes or large fishes when young; all in Arabic; Mosul bleak].

Systematics

Leuciscus maxillaris Valenciennes, 1844 from "rivières de Perse", probably Alburnus capito Heckel, 1843 from "Gebirgsflüssen Kurdistans" (mountain streams of Kurdistan in Heckel (1843b) or "Gebirgsbache in Kurdistan" in Heckel (1847a)), Alburnus Iblis Heckel, 1847 described from the "Gegend um Persepolis oder den Gewässern des Araxes" (= probably the Pulvar (= Sivan) River near Persepolis and the Kor River, both in Fars), Alburnus Schejtan Heckel, 1847 described from the "Araxes bei Persepolis", Alburnus caudimacula Heckel, 1847 described from the "Flusse Kara-Agatsch und bei dem Dorfe Geré (= Qarah Aqaj or Mand River, Fars; Jereh or Jireh is at 29°15'N, 51°58'E according to Edmondson and Lack (2006) and is in the Dalaki River basin (see map in Hohenacker (1845)), and Alburnus megacephalus Heckel, 1847 described from the "Araxes" are synonyms (e.g. according to Berg (1949)). The type locality of Alburnus mossulensis is the "Tigris bei Mossul" according to Heckel (1843b).

Saadati (1977) considers Alburnus caudimacula to be a distinct species found in the Mand River of Fars based on head length being longer (but the ranges overlap) and a shorter scaleless keel (which is individually variable in these fishes according to my observations).

A subspecies, Alburnus mossulensis delineatus Battalgil, 1942, is reported from Diyarbakir on the Tigris River in Turkey.

A hybrid with Acanthobrama marmid was reported from the Hawr al Hammar in southern Iraq by Krupp et al. (1992) who also note that A. mossulensis is probably a synonym of Alburnus sellal Heckel, 1843, a species originally described the Quwayq River at Aleppo. However, they retained mossulensis as a distinct species because of colour differences and the difficulty of obtaining fresh material of sellal in its polluted habitat at Aleppo in Syria (see Vesiland (1993) for habitat photograph). Heckel (1847a) differentiates mossulensis from sellal by the former being more slender and elongate, the pelvic, dorsal and anal fins are more anterior so the caudal peduncle is more elongate, the eyes are larger and lower on the head, and there is a lead-coloured stripe separating the upper third of the body from the lower part. Berg (1949) considers that A. mossulensis may be nothing more than a subspecies of A. sellal. A Principal Components Analysis on the types of mossulensis and sellal using 32 morphometric and meristic characters showed some separation between the two taxa and a Discriminant Function Analysis separated most, but not all, specimens. The evidence is not conclusive for separation or synonymy and the taxa are left as distinct in this work. A. mossulensis itself may be a species complex with taxa in isolated basins in Iran distinct. There are names available for some of these (see synonyms above).

If mossulensis is a synonym of sellal, then the nominal taxa Alburnus hebes Heckel, 1843, Alburnus microlepis Heckel, 1843 and Alburnus pallidus Heckel, 1843, all from the Kueik (= Quwayq) River at Aleppo (Heckel, 1843b), would have to be added to the synonymy of sellal as indicated by Berg (1949), Krupp (1985c) and Eschmeyer's "Catalog of Fishes" (downloaded 1 September 2007). The 3 syntypes of Alburnus hebes seen by me in the Naturhistorisches Museum Wien were 58.8-156.5 mm standard length (NMW 17558-17560) (but Eschmeyer et al. (1996) list NMW 55523 for these syntypes, and the card index had this number in 1997; possibly they were renumbered). One of these fish is designated as the lectotype. The holotype of Alburnus microlepis (NMW 55655) measures 119 mm standard length (Krupp, 1985c). The holotype of Alburnus pallidus (NMW 55720) measured 76.6 mm standard length.

Krupp (1985c) gives details on the syntypes of Alburnus sellal held at the Naturhistorisches Museum Wien. Six syntypes of A. sellal, 124-140 mm standard length, are under NMW 55665 (2 fish, 137.2-141.3 mm standard length, my measurements) and NMW 55666 (4, 126.9-142.7 mm standard length), and 3, 110-152 mm standard length, are under NMW 55664 (1, 110.5 mm standard length) and 55667 (2, one of which is designated as the lectotype, 140.7-155.4 mm standard length). Eschmeyer et al. (1996) list NMW 55664-67 as having 1, 2, 4, and 2 fish in each number in the series and also 2 syntypes (RMNH 2666) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW. The catalogue in Vienna lists 8 specimens of A. sellal.

The syntypes of A. mossulensis are under NMW 55656 (2 fish, 111.2-118.4 mm standard length, my measurements), NMW 55717 (2, 83.0-89.4 mm standard length), and NMW 55718 (2, 101.9-131.5 mm standard length). Two syntypes of Alburnus mossulensis are in the Senckenberg Museum Frankfurt (SMF 402, formerly NMW) (F. Krupp, pers. comm., 1985; 80.1-102.7 mm standard length). Eschmeyer et al. (1996) also list NMW 77723 (2, 90.4-135.4 mm standard length) and 1 possible syntype in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2644). The catalogue in Vienna lists 6 specimens of A. mossulensis, with one specimen from NMW 77723 as the lectotype.

Seven syntypes of Alburnus iblis are in the Naturhistorisches Museum Wien under NMW 55524 and measure 91-165 mm standard length (Kähsbauer, 1964; 92.9-172.3 mm standard length by my measurements). One of these fish is designated as the lectotype. The catalogue in Vienna lists 8 specimens in one column and 38 in the adjacent column.

Two syntypes of Alburnus megacephalus are under NMW 55627 and measure 160-162 mm standard length (Kähsbauer, 1964; 162.9-166.1 mm standard length by my measurements); 2 specimens are listed in the Vienna catalogue. One of these fish is the lectotype.

Fifteen syntypes of Alburnus caudimacula are under NMW 55506 and measure 38.5-118.4 mm standard length; the catalogue in Vienna lists 8 specimens in one column and what appears to be 26 specimens in the adjacent column although this may be 20 fish with 6 set aside for A. schejtan. The Rijksmuseum van Natuurlijke Historie, Leiden has 4 syntypes under RMNH 2654, formerly in NMW (Eschmeyer et al., 1996).

Five syntypes of Alburnus capito measure 48.7-101.9 mm standard length (NMW 55505) although the catalogue in Vienna only lists 4 fish.

Four syntypes of Alburnus schejtan measure 71.7-112.6 mm standard length (NMW 22281) and one of these is designated as the lectotype, 2 syntypes measure 104.5-112.3 mm standard length (NMW 55663), 2 syntypes measure 91.8-100.0 mm standard length (NMW 55719), and 2 syntypes measure 81.6-94.4 mm standard length (NMW 55721).

Two syntypes of Leuciscus maxillaris, 165-166 mm total length, are stored in the Muséum national d'Histoire naturelle, Paris (as 13954 according to Fang (1942) or as A.3954 according to Bertin and Estève (1948), M. L. Bauchot, in litt., 1982, and my observations). Fang (1942) regards maxillaris as a distinct species in Alburnus. My measurements were 136.7-136.9 mm standard length.

Krupp (1985c) refers 5 specimens from the type series of Alburnus doriae to his Alburnus sellal and 2 specimens to Leuciscus (= Squalius) lepidus.

Bianco and Banarescu (1982) felt that their samples showed clinal variation from northwest to southeast, with numbers of anal fin branched rays, lateral line scales and gill rakers gradually decreasing. Their fish from the upper Tigris River basin in Turkey not far from Mosul (the type locality) and from the Pulvar River (Kor River basin of Fars) form one subspecies while those from the Mand and Kul River basin draining to the Persian Gulf in Fars are a distinct subspecies. Available names for the former subspecies include capito, iblis, schejtan and megacephalus, the latter requires a new name according to Bianco and Banarescu (1982). The Tigris-Kor sample could be A. mossulensis mossulensis and the Mand-Kul sample A. mossulensis caudimacula (see above). However, Bianco and Banarescu (1982) are correct to point out that variation in this species has not been fully examined, local environmental conditions such as temperature can affect scale counts and the problem of the relationship of A. sellal remains to be resolved. They found in 7 specimens of sellal that scale counts at 71-77 (in contrast to 66-70 in Berg (1949)) overlapped with mossulensis counts. Berg's (1949) and my counts are very wide for A. mossulensis, suggesting that local environment may govern meristic characters as widely demonstrated for fishes. Subspecies recognition requires much further work as Bianco and Banarescu (1982) acknowledge by not proposing a new name for the Mand-Kul fish.

Key characters

The short, naked ventral keel, usually 8 branched dorsal fin rays, distribution, and the characters in the table under A. atropatenae can be used to identify this species.

Morphology

Dorsal fin with 3 unbranched and 7-9 branched rays, anal fin with 3 unbranched and 10-14 branched rays. Pectoral fin branched rays 13-18, pelvic fin branched rays 7-9. Lateral line scales 58-89. Gill rakers 11-18. Pharyngeal teeth 2,5-4,2, with hooked tips and serrated edges to the crowns. Variants include 2,5-5,2, 3,5-5,3 and 2,5-5,3. Populations vary sympatrically in total vertebral counts: 40-43 and 42-45; and in abdominal counts 20-22 and 22-24 (Bogutskaya et al., 2000). The karyotype of fish from the Kızılırmak River in Turkey was 2n=48 (Gül et al., 2000) but this species does not occur in this area.

Meristics for Iranian specimens:- branched dorsal fin rays 7(16), 8(303) or 9(13); branched anal fin rays 10(20), 11(200), 12(104) or 13(8); pectoral fin branched rays 13(2), 14(30), 15(110), 16(134), 17(50) or 18(6), branched pelvic fin rays 7(30), 8(288) or 9(18); lateral line scales 58(1), 59(-), 60(2), 61(1), 62(5), 63(3), 64(9), 65(8), 66(9), 67(12), 68(11), 69(8), 70(18), 71(17), 72(26), 73(21), 74(28), 75(20), 76(19), 77(26), 78(28), 79(21), 80(15), 81(10), 82(5), 83(1), 84(2), 85(2), 86(2), 87(1), 88(1), or 89(1); and total gill rakers 11(7), 12(54), 13(111), 14(101), 15(46), 16(11), 17(1) or 18(1).

Sexual dimorphism

Unknown but males do develop tubercles in the breeding season.

Colour

Overall colour is silvery. The back is a bluish- or reddish-brown, bluish-black or blackish. A dark, lead-coloured stripe runs along and above the mid-flank and has a width about the same as the eye diameter. The stripe may only be evident posteriorly. Scales above the lateral line have fine melanophores at their base. Lateral line scales can have pigment spots above and below the tube near the base of each scale, but this is not as marked as insome Alburnoides spp. The dorsal, anal and caudal fins are margined with black, the latter the darkest. There may be a black spot at the caudal fin base and the first pectoral fin ray may be black dorsally. The pectoral, pelvic and anal fins are yellowish at their base. Pelvic and anal fins may be reddish. The peritoneum is brown but may be thickly speckled with black-brown spots and thus appear almost black.

Size

Reaches about 22 cm (Ergene, 1993).

Distribution

Found in the Tigris-Euphrates basin and adjacent basins. In Iran it is recorded from the Tigris River, Gulf, Lake Maharlu, Kor River and upper reaches of the Hormuz basins (M. Hafezieh, pers. comm.; Berg, 1949; Bianco and Banarescu, 1982; Abdoli, 2000) and questionably from the Esfahan basin (Abdoli, 2000). Records also include the Shapur and Dalaki rivers in the Gulf basin (Gh. Izadpanahi, pers. comm., 1995) and the upper Mand including Qara Agaj reach and Shur tributary, Shur tributary to Dasht-e Palang; upper Zohreh, Marun and Jarrahi, upper Karun and Khersan, Dez, whole middle to upper Karkheh basin (Simarreh, Qarasu, Gav Masiab) (Abdoli, 2000).

Zoogeography

Its former position in the genus Chalcalburnus indicates a relationship with fishes occurring in the Black-Caspian seas basin.

Habitat

This species is found in streams, rivers, lakes, reservoirs and marshes. Al-Habbib (1981) has demonstrated experimentally for specimens taken from the Aloka River, north of Mosul, Iraq that this species can survive temperatures in the range of about 1.25-36.2°C when acclimated (fish were identified incorrectly as Chalcalburnus chalcoides). Epler et al. (2001) found it to be the second most dominant species of fish (identified as A. sheitan) in lakes Habbaniyah, Tharthar and Razzazah in Iraq, comprising 10% of all fish collected. This was one of the most abundant species in the recovering marshes of southern Iraq in 2005-2006 (Hussain et al., 2006). Abbasi et al. (2009) in their study of wetlands in Hamadan Province found this species was dominant out of 23 species at 28%.

Age and growth

Jawad (2004) used eye lens diameter for ageing the young (up to age 3) of this species from the marshes north of Basrah. Ergene (1993) studied the growth of this species in the Karasu of Turkey and found 4 age groups, and mentions 5 age groups for another Turkish study. Mean fork length is 118.2 mm, 131.0 mm, 145.2 mm and 163.3 mm respectively. Condition factors for these age groups were 0.87, 0.85, 0.84 and 0.86. Türkmen and Akyurt (2000) also working on this species in the Karasu River found age groups 1 to 6 with age group 3 the most abundant. The mean condition factor for males and females was 1.023 and 1.047 respectively. Age-length, age-weight (von Bertalanffy equations) and length-weight relationships were also calculated as lt = 20.41[1-e-0.2485 (t+1.47)], lt = 21.59[1-e-0.1978 (t+2.13)], W = 80.77 (1-e-0.2485 (t+1.47)2.828, W = 103.63 (1-e-0.1978 (t+2.13)3.082, LogW = -1.796 + 2.828 LogFL (r = 0.943) and LogW = -2.097 + 3.082 LogFL (r = 0.946) respectively. Length and age at first maturity were 1.26 years and 9.24 cm for males and 1.81 years and 9.65 cm for females in the Karasu River, Turkey; age group 7 was the oldest recorded (Yıldırım et al., 2007).

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 76 Iranian fish measuring 3.15-8.14 cm standard length. The a-value was 0.0197 and the b-value 2.903 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Younis et al. (2001b) found Shatt al Arab, Iraq fish feeding on phytoplankton (algae and diatoms) at 44%, followed by organic detritus at 36.7% (33% in a table), and arthropods at 3.1%, It had a dietary overlap of 89% with Barbus (= Carasobarbus) luteus in May, the highest in the study. In a study of the recovering Hammar Marsh, Iraq diet was 67.95% insects and 14.34% algae with diatoms, plants, crustaceans and fish at less than 10% each, in the Hawr al Hawizah 66.2% insects and 19.2% algae, with amounts of diatoms and crustaceans being less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 73.7% insects and 13.1% algae with diatoms, plants and crustaceans at less than 10% each (Hussain et al., 2006).

Reproduction

Berg (1949) reports a female 15.5 cm long with mature eggs. Qarmat Ali River, Iraq fish had a fecundity of 1926-11,779 eggs (Saud, 1997). Yıldırım et al. (2007) examined this species in the Karasu River of Turkey and found a male:female sex ratio of 1:1.08, not significantly different from 1:1, a fecundity range of 3012 to 11,427 eggs, significant correlations between fecundity and fork length, total weight, age and gonad weight, and a spawning season from June to August when water temperature attained 15ºC.

Parasites and predators

Molnár and Jalali (1992) describe a new species of monogenean, Dactylogyrus holciki, from this species in the Beshar River of the Persian Gulf drainage. Gussev et al. (1993b) report the monogenean, Dactylogyrus chalcalburni, from Alburnus alburnus in the Zayandeh Rud but this fish does not occur there. The parasite may have been found in Alburnus mossulensis. González-Solís et al. (1997) report Rhabdochona denudata, Contracaecum sp. larvae and Proleptinae larvae (Nematoda) from this species in the drainage of Lake Maharlu and Contracaecum sp. larvae in the drainage of the Kor River, both in Fars. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from the Beshar River of the Tigris basin in a Chalcalburnus sp., presumably this species. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea and Diplostomum spathaceum. Barzegar et al. (2008) also record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species.

Economic importance

This species has been used in the preparation of fish meal in Iraq.

Conservation

An abundant species where studied, it appears to be under no threat in Iran. Endangered in Turkey (Fricke et al., 2007).

Further work

Its taxonomic status in relation to its Levant relative remains unresolved and the relation between lowland and mountain populations in Iran needs careful analysis. Its biology in Iran has yet to be studied in detail.

Sources

Type material: See above, Alburnus capito (NMW 55505), Alburnus caudimacula (NMW 55506), Alburnus hebes ((NMW 17558-17560 or NMW 55523), Alburnus iblis (NMW 55524), Alburnus megacephalus (NMW 55627), Alburnus microlepis (NMW 55655), Alburnus mossulensis (NMW 55656, 55717, 55718, 77723, SMF 402), Alburnus schejtan (NMW 22281, NMW 55663, 55719, 55721), Alburnus sellal (NMW 55664, 55665, 55666, 55667), Leuciscus maxillaris (MNHN A.3954).

Iranian material: CMNFI 1977-0510A, 44, 35.7-154.6 mm standard length, Fars, Pulvar River tributary (29º59'30"N, 52º54'E); CMNFI 1979-0025, 87, 19.1-138.2 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0027, 24, 59.8-105.0 mm standard length, Fars, Chehel Cheshmeh (ca. 29º43'N, ca. 52º04'E); CMNFI 1979-0028, 55, 19.1-122.6 mm standard length, Fars, Kor River drainage (no other locality data); CMNFI 1979-0036, 22, 82.3-115.1 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0047, 7, 41.4-78.2 mm standard length, Fars, Ab-e Paravan spring (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0054, 17, 39.8-95.6 mm standard length, Fars, Shur River tributary (28-29º58-03'N, 52º34-35'E); CMNFI 1979-0061, 51, 32.9-131.1 mm standard length, Fars, Pulvar River tributary (30º04'N, 53º01'E); CMNFI 1979-0067, 55, 11.1-107.9 mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0070, 44, 35.0-98.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0071, 12, 65.3-104.3 mm standard length, Fars, qanat 23 km from Pol-e Khan (ca. 30º00'N, ca. 52º38'E); CMNFI 1979-0073, 26, 50.0-93.3 mm standard length, Fars, Mand River (ca. 29º42'30"N, ca. 52º01'30'E); CMNFI 1979-0074, 39, 23.8-94.0 mm standard length, Fars. Mand River (29º41'N, 52º06'E); CMNFI 1979-0117, 16, 33.4-130.0 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0128, 9, 43.2-102.5 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0154B, 1, 46.9 mm standard length, Fars, stream at Koorsiah village (28º45'30"N, 54º24'E); CMNFI 1979-0155, 2, 56.2-64.7 mm standard length, Fars, spring at Gavanoo village (28º47'N, 54º22'E); CMNFI 1979-0156, 11, 49.0-74.4 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 53, 31.8-86.6 mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, 13, 73.5-108.9 mm standard length, Fars, qanat between Now Bandegan and Qaziabad (28º54'N, 53º53'30"E); CMNFI 1979-0160, 22, 32.4-106.0 mm standard length, Fars, spring at Arteshkhadeh Pomp (29º09'N, 53º37'E); CMNFI 1979-0272, 11, 40.5-130.0 mm standard length, Lorestan, river at Nokhor (ca. 33º40-47'N, ca. 48º28-45'E); CMNFI 1979-0278, 4, 75.5-88.2 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 3, 68.7-91.4 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0282, 19, 40.2-131.3 mm standard length, Lorestan, river at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0284, 30, 73.1-98.3 mm standard length, Kermanshahan, Qareh Su drainage (34º16'N, 46º48'30"E); CMNFI 1979-0285, 4, 124.7-136.8 mm standard length, Kermanshahan, Qareh Su drainage (34º26'N, 46º37'E); CMNFI 1979-0289, 2, 125.3-142.1 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 4, 146.9-171.4 mm standard length, Kermnanshahan, Diyala River drainage (34º31'N, 45º35'E); CMNFI 1979-0348, 4, 68.0-78.8 mm standard length, Fars, 2 km from Pol-e Berengie (ca. 29º28'N, ca. 52º32'E); CMNFI 1979-0352, 2, 88.5-93.9 mm standard length, Khuzestan, Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0499, 3, 104.8-133.6 mm standard length, Fars, irrigation ditch on road to Dariush Dam (30º04'30"N, 52º36'E); CMNFI 1979-0500, 2, 112.2-116.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E).

Comparative material: BM(NH) 1981.4.13:9-11, 3, 64.3-72.8 mm standard length, Aloka River near Mosul (no other locality data); CMNFI 1980-815, 2, 88.9-107.2 mm; CMNFI 1980-1036, 2, 11.6-145.8 mm standard length, Turkey, ?check this sample for gill raker count ;

Alburnus zagrosensis
Coad, 2009


Holotype

Male, Morghab, courtesy of Jörg Freyhof
Male, Morghab, courtesy of Jörg Freyhof

Common names

None.

Systematics

The holotype (CMNFI 1979-0248) is a male 81.6 mm SL from Chahar Mahall va Bakhtiari, stream, 3 km east of Boldaji, upper Karun River basin (31°55’N, 51°05’E); paratypes are CMNFI 1979-0248A, 128 ♂ 48.4-89.5 mm SL same locality as holotype and CMNFI 1979-0246, 49, 52.5-92.6 mm SL, Chahar Mahall va Bakhtiari, stream, 8 km west of Boldaji, upper Karun River basin (31°57’30”N, 50°59’E). The species is named for the Zagros Mountains of western Iran where it was collected at altitudes over 2300 m.

Key characters

This species is distinguished from other Iranian and Tigris-Euphrates basin Alburnus by such characters as high lateral line scale count (67-83; 64 or lower in alburnus, atropatenae, caeruleus, doriae, filippii), low anal fin branched ray count (9-10; usually 12 or higher in alburnus, caeruleus, chalcoides), and low total gill raker count (12-14; 16 or higher in alburnus, chalcoides). Other characters include a ventral keel almost absent to almost complete, a high frequency of 7 branched dorsal fin rays (35.3%), a high frequency of 7 branched pelvic fin rays (62.7%), a total vertebral count mode at 42, absence of a prominent mid-flank stripe, and small size (females mature at 88.5 mm SL).

Morphology

The body is elongate and a vertical oval in cross-section, somewhat compressed but not deep. The upper and lower profiles are a gentle arch. The snout is short and pointed. The mouth is slightly superior, almost terminal, oblique, and extends back to a level with the mid-nostril. The mouth tip is on a level with the upper half of the eye. The anal fin origin is posterior to the dorsal fin insertion. The dorsal fin margin is rounded and the anal fin margin is straight. The lateral line is decurved and is on the midline of the body only on the posterior half of the caudal peduncle. There is a pelvic axillary scale. The naked ventral keel is flanked for six scale rows anterior to the anal fin.

Dorsal fin with 3 unbranched rays and 7(18) or 8(33) branched rays, anal fin with 3 unbranched rays and 9(29) or 10(22) branched rays, pectoral fin with 14(5), 15(25), 16(15) or 17(6) branched rays, pelvic fin with 7(32) or 8(19) branched rays, lateral line scales to hypural fold 67(1), 69(2), 70(3), 71(2), 72(6), 73(6), 74(2), 75(4), 76(6), 77(7), 78(3), 79(3), 80(2), 81(3), or 83(1), scales around caudal peduncle 20(1), 21(10), 22(18), 23(11), 24(10) or 25(1), predorsal scales 32(1), 33(5), 34(1), 35(8), 36(8), 37(5), 38(9), 39(3), 40(6), 41(4) or 42(1), scales between lateral line and dorsal fin origin 12(1), 13(2), 14(22), 15(17), 16(8), or 18(1), scales between lateral line and anal fin origin 5(6), 6(37) or 7(8), scales between lateral line and pelvic fin origin 4(6), 6(25), 6(15) or 7(5), total gill rakers 12(20), 13(19) or 14(12), short and usually reach past the anterior base of the adjacent raker when appressed, abdominal vertebrae 21(14), 22(34) or 23(3), caudal vertebrae 19(17), 20(32) or 21(2), and total vertebrae 40(4), 41(19), 42(27) or 43(1).

From 0 to 5 scales at the end of the lateral line are not pored. The belly is rounded and has a naked keel in front of the anal fin, varying in extent. The naked keel may be flanked by one to 11 scales, or from almost no keel to a keel extending from the anal fin almost to the pelvic fins base. Scales from below the dorsal fin have fine but not numerous circuli, a subcentral anterior focus and few radii on the anterior and posterior fields with no radii on the lateral fields. Pharyngeal teeth number 2,5-4,2 (6), 2,5-4,1(2) or 1,5-4,2(2). Teeth are hooked at the tip and have serrated edges to the narrow grinding surface below the tip. The gut is an elongate S-shape with a small anterior loop to the left.

Sexual dimorphism

Small tubercles are present on the mature male, on the sides and top of the head, on the dentaries, as fine ones lining the scale margins, most evident on caudal peduncle scales, and on the upper pectoral fin rays. Significant differences between males and females were found in interorbital width (wider in males), caudal peduncle depth (deeper in males), and dorsal, anal, pectoral and pelvic fin lengths (longer in males).

Colour

Fins are mostly immaculate with only slight traces of melanophores. The body is darker dorsally and the upper flank is much darker than the lower flank with an abrupt transition in pigmentation between the two halves along the midline. The anterior flank has a less defined transition with some thin bars extending downward for a short distance. The mid-flank dark pigmentation forms a weak stripe, well-defined by its ventral edge above the anal fin level but the dorsal edge is less well-defined and merges with the upper flank pigmentation. The end of the caudal peduncle has a broad, fan-shaped pigmentation. A broad stripe is present predorsally and postdorsally. The predorsal stripe may be separated into two thin stripes, one on each side of the mid-line with a very thin stripe between them. After 32 years in preservative, this pigment pattern is less evident and the division between the epaxial and hypaxial muscle masses is marked by an apparent thin line of dark pigment as the most prominent feature. The peritoneum is silvery with numerous scattered melanophores grading to fish with continuous melanophores and dark brown overall.

Size

Attains 92.6 mm standard length.

Distribution

Known only from the type locality in the upper Karun River basin.

Empty square = holotype, black square = paratypes, of A. zagrosensis; black circles = A. mossulensis

Empty square = holotype, black square = paratypes, of A. zagrosensis; black circles = A. mossulensis

Zoogeography

This species is related to A. mossulensis which is found in same basin (Tigris-Euphrates) and may have evolved from an isolated population in the upper Karun River.

Habitat

The type locality was at an altitude of 2360 m, water temperature was 22°C at 1545 hours, pH was 6.2 and conductivity 0.45 mS. The stream width was 3-4 m, depth 1 m maximum, the bottom was muddy and the water cloudy. Current was slow, the shore grassy and there was a moderate amount of aquatic vegetation, mostly Myriophyllum. Other species caught were the cyprinid Chondrostoma regium and the tooth-carp Aphanius vladykovi.

The second locality was at an altitude of 2380 m, water temperature was 25°C at 1450 hours, pH was 6.2 and conductivity 0.4 mS. The stream width was 1-2 m, depth 1 m maximum, the bottom was stones and mud and the water muddy. Current was fast, the shore grassy and there was some aquatic vegetation. Some of this material was caught while leaping up a raceway. Other species caught were the cyprinids Chondrostoma regium and a Capoeta sp.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

Unknown.

Economic importance

None.

Conservation

Although known from only two localities in the upper Karun River basin of Iran, this species may not be threatened other than by water abstraction and pollution.

Further work

Distribution, conservation and biology of this species needs investigation.

Sources

See Coad (2009).

Type material: See above.

Genus Aspidoparia
Heckel, 1847

This Oriental genus has only 2 species, one of which enters southeastern Iran. Mirza (2000) proposes that the members of the genus Aspidoparia be placed in a new subfamily, Aspidoparinae. Cabdio Hamilton, 1822 is presumably a senior synonym of Aspidoparia but has not been extensively used in literature on these species so Aspidoparia is retained here.

It is characterised by an elongate and almost cylindrical body with a rounded abdomen, the head has a broad ring of suborbital bones, the mouth is small and inferior, the roof of the mouth has a papillose nodule, the lower jaw has a sharp, crescentic bony edge, no barbels, pharyngeal teeth in 2-3 rows, dorsal fin short, anal fins short to moderate, scales moderate in size, lateral line decurved and running on the lower half of the caudal peduncle, and the gut is long and coiled.

Aspidoparia morar
(Hamilton, 1822)

Common names

None.

[waspi or common chilwa in Pakistan].

Systematics

No relevant synonyms. This species was originally described from the Yamuna and Tista rivers, India. No types are known (Eschmeyer et al., 1996).

Key characters

The suborbital ring of bones is large and distinctive, being almost as deep as the eye, and this feature is unique in southeastern Iranian cyprinids.

Morphology

The snout is short and rounded and overlaps the upper lip. The mouth is small, ventral and transverse. The lower jaw is straight with a slightly horny cutting edge and no lip. The dorsal fin origin is over or slightly behind the pelvic fin origin. The dorsal fin margin is straight to very slightly emarginate and the anal fin is emarginate.

Dorsal fin unbranched rays 2-3 (the first unbranched ray is very small, usually 3 rays are present but not discernible) and branched rays 6-8, anal fin unbranched rays 2 and branched rays 8-10, pectoral branched rays 9-16, usually 12 or more, and pelvic fin branched rays 7-8. Lateral line scales 36-45. Scales have few anterior and more numerous but not many posterior radii. There is a pelvic axillary scale and several elongate and overlapping scales in the pectoral axil. Gill rakers are very short, not touching the adjacent one when appressed, difficult to count at the fleshy ends of each arch, and numbering about 17-25. Pharyngeal teeth 2,4,5-5,4,2 in the literature but the main row count of 4 teeth observed here differs. The main row teeth have large, oval to oblong flattened crowns. The gut is a very elongate s-shape with a small anterior loop. Total vertebrae 36-37. The chromosome number is 2n=48 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 6(1) or 7(18); anal fin branched rays 8(4) or 9(15); pectoral fin branched rays 9(1), 12(5), 13(11) or 14(2); pelvic fin branched rays 7(19); lateral line scales 36(1), 37(4), 38(1), 39(5), 40(4), 41(2), 42(1) or 45(1); scales above the lateral line 7(10) or 8(9); scales below the lateral line to the anal fin 3(1), 4(12) or 5(6); scales between the lateral line and the pelvic fin 4(13), 5(5) or 6(1); predorsal scales 17(1), 18(3), 19(2), 20(2), 21(3), 22(3), 23(2) or 24(2); caudal peduncle scales 15(1), 16(5), 17(6), 18(5), 19(1); total gill rakers 17(1), 18(2), 19(2), 21(2), 22(4), 23(2) or 25(1); pharyngeal tooth count 2,4,4-4,4,2(5), 2,4,4-4,4,1(1), or 2,3,4-4,4,2(1), and total vertebrae 36(3) or 37(9).

Sexual dimorphism

Unknown.

Colour

Back light brown to brown-green with the flanks very silvery to silvery-yellow and the belly lighter. There is a golden stripe along the flank. Fins are a distinct dark yellow. The caudal fin may be yellow to orange and paired fins a very light orange-yellow. Preserved fish have immaculate fins except for the caudal fin which has some melanophores lining the rays, a broad stripe along the midline of the back, and fine melanophores on the back and upper flank. Some fish have small, dark dots on the back and upper flank. The peritoneum is black.

Size

Attains 20 cm (Malhotra and Munshi, 1985).

Distribution

This species is reported from the Makran and Mashkid River basins in Pakistan (Mirza, 1992) and eastwards to Thailand. The Iranian distribution encompasses the Mashkel (= Hamun-e Mashkid) and Makran basins, the latter westwards to the Straits of Hormuz (Kiabi and Abdoli, 2000).

Zoogeography

The species and genus reaches its westernmost limit of distribution in southeastern Iran. Barriers to further dispersal are unknown but it may be limited by temperature, habitat availability and poor recent connections between streams in the Makran and the southern deserts of Iran.

Habitat

This species favours streams with slow current.

Age and growth

A female, 9.8 cm total length, from Iran had mature eggs (Berg, 1949).

Food

This minnow is a carni-omnivore and a voracious feeder (Bhattacharjee and Dasgupta, 1988). Iranian specimens contained no discernible food items in their guts.

Reproduction

Spawning occurs from February to April in India (Malhotra and Munshi, 1985), Iranian specimens caught in December were not mature suggesting a later spawning season.

Parasites and predators

Jalali et al. (2000) describe two new species of monogeneans, Dactylogyrus yousefpouri and D. mobedii, from this species in the Bahu Kalat River of Baluchestan.

Economic importance

Not of any economic importance in Iran but it is eaten in India.

Conservation

Although known from only a few localities in southeastern Iran, this species may not be threatened other than by water abstraction and pollution.

Further work

The biology of this species, which is at its westernmost range limit in Iran, is unknown. There are some minor differences in characters with literature reports, particularly in pharyngeal tooth count, but sample sizes do not permit an adequate comparison for this wide-ranging species.

Sources

Iranian material: CMNFI 1979-0316, 1, 22.1 mm standard length, Baluchestan, stream in Sarbaz River drainage (26º48'N, 61º02'E);CMNFI 1979-0322, 7, 42.3-86.3 mm standard length, Baluchestan, Dashtiari River (ca. 25º45'N, ca. 61º26'E); CMNFI 1979-0333, 7, 17.7-69.5 mm standard length, Baluchestan, Mashkid River (ca. 27º05'N, ca. 63º12'E); CMNFI 1979-0334, 10, 22.8-62.0 mm standard length, Baluchestan, Mashkid River (27º04'N, 62º54'E): OSU 8123, 5, 45.7-50.6 mm standard length, Baluchestan, Srabaz River (no other locality data).

Comparative material: BC55-61, 2, 67.0-68.2 mm standard length, India, Barakar River near Tillya Dam (no other locality data).

Genus Aspiolucius
Berg, 1907

Aspiolocius esocinus
(Kessler, 1874)

Recorded from the Karakum Canal in Turkmenistan (Sal'nikov, 1995) and may eventually be found in the Tedzhen (= Hari) River basin of Iran. No Iranian record.

Genus Aspius
Agassiz, 1832

The asps comprise 2 species found in Europe and Southwest Asia. Both species are found in Iran.

This genus is characterised by an elongate, rounded and large body, small scales, a large mouth with the lower jaw projecting, lower jaw with a tubercle fitting into a notch in the upper jaw, no barbels, pharyngeal teeth in 2 rows, pointed and hooked, gill rakers short and wide apart, short dorsal fin without a thickened ray, a long anal fin, a scaled keel behind the pelvic fins, and gill slits very wide such that the branchiostegal membranes attach under the posterior end of the eye. Perea et al. (2010) using mitochondrial and nuclear DNA propose the synonymy of this genus with Leuciscus Cuvier, 1816.

Aspius aspius
(Linnaeus, 1758)

Anzali Shore, November 2010, courtesy of K. Abbasi
Anzali Shore, November 2010, courtesy of K. Abbasi

Common names

ماش ماهي (= mash mahi, not apparently meaning pea fish as the Farsi could indicate), khasham.

[hasam or khasham in Azerbaijan; krasnogubyi zherekh or redlip asp in Russian; Caspian asp, South Caspian asp].

Systematics

Cyprinus Aspius was described originally from lakes of Sweden.

Cyprinus Rapax Leske, 1774 described from Leipzig, Germany, Cyprinus taeniatus Eichwald, 1831 described from the Kura River at Mingechaur, Aspius erytrostomus Kessler, 1877 (sic, sometimes spelt erythrostomus or erithrostomus) described in part from the Caspian Sea and Kura River, Azerbaijan and from the Aral Sea and lower part of the Amu Darya, Uzbekistan, and Aspius transcaucasicus Warpakhovskii, 1895 from the Lenkoran River and Lake Bussadagny, Azerbaijan, are synonyms. Aspius aspius taeniatus (Eichwald, 1831) is the subspecies found in the Caspian Sea.

The types of Cyprinus aspius described from Swedish lakes are unknown (Eschmeyer et al., 1996).

Eschmeyer et al. (1996) give Aspius transcaucasicus Varpakhovskii, 1896, although Berg (1948-1949) gives 1895; possibly the volume year is 1895 but the work did not appear until 1896. Varpakhovskii is a variant spelling in transliteration from the Russian. Syntypes of this synonym are in the Zoological Institute, St. Petersburg under ZISP 10488 (2) and ZISP 10497-48 (sic, in Eschmeyer et al. (1996) but should read 10497-98 with 5 and 2 specimens respectively (Kottelat, 1997)).

Rezaei et al. (2012) examined fish from two fishing areas (Tonekabon and Sari) using meristics and morphometrics and found that the populations were distinct but with a relatively high degree of overlap.

Key characters

The subspecies of the southern Caspian Sea is distinguished from the type subspecies of Europe and the northern Caspian Sea since the latter has lower lateral line scale counts of 64-76 as opposed to 62-105, lips never bright red, anal fin branched rays usually 13 instead of 12 (but see Iranian fish below), and height of dorsal fin usually longer than distance from snout tip to posterior edge of preopercle. Characters of the genus and distribution serve to separate it from other cyprinids in Iran.

Morphology

Dorsal fin branched rays 7-10, usually 8, after 2-3, usually 3, unbranched rays, and anal fin branched rays 11-16, usually 12 (but see below), after 3-4, usually 3, unbranched rays, pectoral fin branched rays 14-17 and pelvic fin rays 7-9. Lateral line scales 62-105. The scales have a central focus, fine circuli and few posterior and anterior radii. There is a pelvic axillary scale. There is a scaled keel behind the pelvic fins. The lower jaw tip projects and fits into a notch in the upper jaw. Gill membranes are narrowly attached to the isthmus, almost under the posterior eye margin. Gill rakers 8-11, very short and club-shaped, almost reaching or not reaching half way to the raker below when appressed. Pharyngeal teeth usually 3,5-5,3, sometimes 2,5-5,3 or with 6 teeth in the main row, teeth elongate, compressed and obviously hooked. Gut an elongate s-shape. Vertebrae 49-51. The chromosome number is 2n=50-52 (Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(6); anal fin branched rays 13(6); pectoral fin branched rays 18(3); pelvic fin branched rays 8(3); lateral line scales 68(1), 72(2), 73(1), 74(1) or 75(2); total gill rakers 8(1) or 9(2); pharyngeal teeth 3,5-5,3(3); and total vertebrae 50(2) or 51(1).

Sexual dimorphism

Unknown.

Colour

The overall colour is silvery with the back a blackish-olive or greenish-grey. The iris is silvery with a narrow golden circle around the pupil and a little grey pigment on the upper half. Lips are silvery with a little grey over the upper one. Both lips and iris are often bright red. The dorsal and caudal fins are grey and the other fins are transparent without pigment. Fins may be tinged reddish. Peritoneum silvery to brown.

Size

Reportedly attains 1.2 m and 20.0 kg, possibly over 30.0 kg (Machacek (1983-2012), downloaded 27 July 2012). The largest of 12,000 fish from the lower Kura River was 77 cm total length, males averaged 61 cm and females 64 cm. The average weight of 105,500 fish caught in 1927-1929 was 2.72 kg, females 2.93 kg (based on 1500 fish), males 2.34 kg and the heaviest fish was 5.5 kg (Berg, 1948-1949).

Distribution

Found from the Rhine and north of the Alps in Europe to the drainages of the Black, Caspian and Aral seas including their southern shores.

This species has been reported from Astara to Gorgan Bay in rivers and marshes and in the Caspian Sea of Iran (Nedoshivin and Iljin, 1929; Derzhavin, 1934; Berg, 1948-1949; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009; Rezaei et al., 2012). Formerly reported from the Anzali Mordab but no longer present (Holčík and Oláh, 1992) although reported from the Siah-Keshim Protected Region of the Anzali Mordab and Anzali Shore by Riazi (1996) and the Anzali Shore by K. Abbasi (see photograph above). Found also in the Aras River Dam (Jolodar and Abdoli, 2004).

Also recorded from the Uzboi lakes, Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually appear in the Tedzhen (= Hari) River basin in Iran.

Zoogeography

The closest relative of this species lies to the south and indicates a connection between Euro-Mediterranean and/or Black-Caspian-Aral seas basins.

Habitat

In the waters of Dagestan, asp begin to migrate upriver in October, peaking at the end of November and the beginning of December. They overwinter in deep holes, emerging in early spring as rivers flood and move to the spawning grounds. These grounds include river channels, open lake areas with substantial flow and only rarely places weakly overgrown with very coarse submerged vegetation such as reeds and rushes. After spawning the asps return to the Caspian Sea (Shikhshabekov, 1979). Knipovich (1921) reports this species from depths of 14.6-16.5 m, and possibly deeper, in the Iranian Caspian Sea. Riazi (1996) reports that this species is native (resident) to the Siah-Keshim Protected Region of the Anzali Mordab.

Age and growth

Life span in the Volga delta is 7-8 years with the bulk of the population mature at 6 years (Ali, 1974). In the waters of Dagestan life span is 8 years with maturity at 4 years. Mature males and females are 41-58 cm long and weigh 840-2800 g (Shikhshabekov, 1979). Growth is more rapid in the Kura River of Azerbaijan than in other rivers in the former Soviet Union. Fish taken from commercial catches in Iran are mostly 3-6 years old, 38.1-56.7 cm long and weigh 631-2241 g (Razivi et al., 1972) or 3-6 years and 33-63 cm total length (Holčík and Oláh, 1992). Growth is rapid in the latter report, fish reaching 1 kg during the fourth year of life. Maximum life span may be 15 years and Rezaei et al. (2012) reported fish up to 10+ years in Iran.

Food

This species is a solitary predator on other fishes such as gobies (Gobiidae) and silversides (Atherinidae), frogs and even ducklings. An Iranian specimen had the remains of a large crustacean in its gut. Young feed on plankton initially but start to take the fry of fishes at 2-3 months. There is little feeding on the spawning migration.

It may catch other fishes by plunging into shoals at the surface and may leap out of the water as a result. Abdoli (2000) reports Scardinius erythrophthalmus, Atherina boyeri and Blicca bjoerkna as food items in Iran. Surface insects are also eaten.

Reproduction

The spawning season in Gilan is mid-February to late March at 10-13°C with an incubation period of 9-10 days (Hoseinie, 1995).

Spawning is non-intermittent and the period is short (10-15 days) in Dagestan (Shikhshabekov, 1979). Fecundity reaches 483,500 eggs in the south Caspian Sea and maximum egg diameter in the Volga delta is 1.7 mm (Ali, 1974). In Hoseinie's (1995) study of artificial propagation of this species in Iran, large or swollen eggs number 117-277 per gram, and egg diameters 2.0-2.2 mm. Absolute fecundity reaches 264,248 eggs. Abdurakhmanov (1962) gives a maximum fecundity of 342,000 eggs and a maximum egg diameter of 2.4 mm for Azerbaijan populations. Females with ripe eggs are found between mid-April and mid-May at water temperatures of 4-12.2°C, optimally 9-11°C. Up to 20% of Volga asp females do not spawn annually. Eggs develop while between or adhering to stones on the river bed. Young migrate downriver from June to August at age 3-4 months and 5-10 cm length.

Parasites and predators

Molnár and Jalali (1992) record the monogenean Dactylogyrus tuba from this species in the Safid Rud. Masoumian et al. (2005) report the protozoan parasite Chilodonella, sp. from this species in the Aras Dam in West Azarbayjan. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Myxobolus dispar. Sattari (2004) records the presence of the nematode, Eustrongylides excisus, in the body cavity. This parasite can damage muscles in commercial species and render them unsuitable for sale. Sattari et al. (2002, 2004, 2005) and Sattari (2004) records the presence of the nematode, Eustrongylides excisus. This parasite can damage muscles in commercial species and render them unsuitable for sale. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, including Argulus foliaceus from this species.

The Caspian seal, Pusa caspica, is a predator (Krylov, 1984).

Economic importance

This fish is taken in Iran as food but comprises only a small portion of the catch. Nevraev (1929) reports catches of 267 to 2429 fish for the period 1914-1915 to 1917-1918 in the Anzali region. Holčík and Oláh (1992) record the catch in the Anzali region for 1969-1970 and 1970-1971 as 45.2 t and 36.1 t respectively, these being 84% and 69% of the total Iranian catch. In 1921-1930 the annual catch in the lower Kura River averaged 249,000 fish and in 1936 for Azerbaijan the catch weighed 8100 centners and numbered 300,000 fish.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use as food and in sport. The flesh is white and tasty but rather tough.

Conservation

Recruitment in this species is low in Iran because water is taken from the summer spawning streams for irrigation purposes. Spawning success is therefore limited. Larvae of spring spawners are lost when they enter irrigation channels and become stranded in fields (Razivi et al., 1972). Holčík and Oláh (1992) consider the decline in this species to be due to indiscriminate catching of sexually immature fish and, in the Anzali Mordab at least, environmental changes. The Pol-e Astaneh Fish Farm has studied propagation of this species (Keivany and Nasrollahzadeh, 1990) and Hoseinie (1995) demonstrates that artificial propagation is possible. It has also been raised to marketable size in ponds through artificial feeding with ground kilka and a rice product (Annual Bulletin 1993-94, Iranian Fisheries Research and Training Organization, Tehran, p. 81-82, 1995). The Shahid Beheshti hatchery on the Safid River breeds this species (Raymakers, 2002).The asp is bred in the Varvarinsk Hatchery and releases up to 1.5 million yearlings are made into the Kura River, with plans for 8-10 million releases (Kosarev and Yablonskaya, 1994).

Lelek (1987) classifies this species as vulnerable to endangered in Europe. Vulnerable in Turkey (Fricke et al., 2007). 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, habitat destruction, limited range (less than 25% of water bodies), absent in other water bodies in Iran, and absent outside the Caspian Sea basin.

Further work

The distribution and abundance of this species in Iranian waters needs investigation as it is sensitive to environmental changes.

Sources

Iranian material: CMNFI 1970-0526, 2, 236.8-246.1 mm standard length, Gilan, Safid River below Astaneh (37º19'N, 49º57'30"E); CMNFI 1980-0494, 1, 319.6 mm standard length, ? Gilan, Caspian Sea basin (no other locality data); ZISP 3917, 1, 402.0 mm standard length, Gilan, Anzali (no other locality data).

Aspius vorax
Heckel, 1843


Hawr al Azim, courtesy of Asghar Mobaraki
 

Common names

shelej, shalaj, sholge, sholgeh.

[shillik, shillig, shiliq, shelej, shalaj; bu aliawi, abu elawi; called "snake" by American soldiers in Iraq because of the name asp being familiar as the snake that killed Cleopatra; kaschschasch (= voracious) from Heckel (1843b); all in Arabic; Tigris asp].

Systematics

The type locality for this species is the "Tigris bei Mossul" according to Heckel (1843b). Krupp (1985c) reports, and I have examined, a syntype held in the Naturhistorisches Museum Wien under NMW 76776, 261.4 mm standard length. The catalogue in Vienna in 1997 also lists NMW 76785 as a type and this specimen is also 261.4 mm standard length. Eschmeyer et al. (1996) lists a dried skin as a syntype under NMW 16527. The catalogue in Vienna lists 4 fish in spirits and 2 fish stuffed.

Banister (1980) suggests that this species may be close to Aspius aspius, perhaps a clinal variant, since the Caspian Sea basin subspecies, A. a. taeniatus (67-90) has scale counts intermediate between European populations of A. aspius (65-74) and A. vorax (93-105) (Banister's figures). However this may be more apparent than real as there is considerable overlap and frequency distributions are not given. There was insufficient material on hand from Iran to investigate this character in more detail.

Key characters

Characters of the genus coupled with distribution serve to identify this species.

Morphology

The head is long and tapers anteriorly. The mouth is oblique and elongate reaching to the anterior half of the eye. The lower jaw projects and has a symphysis knob fitting into an upper jaw notch. There is a hump as the back rises abruptly after the head. The gill opening is large and extends forward to the posterior eye margin level. Fins are more falcate than in the line illustration when partially collapsed.

Dorsal fin with 2-3 unbranched and 7-9, usually 8, branched rays. Al-Nasiri et al. (1975) give a range of 8-11 (probably 7-10 using my system of counting) dorsal fin rays with a strong mode at 9 (i.e. 8) for 271 fish taken from the Basrah fish market from January to June. Anal fin with 2-3 unbranched and 9-13 branched rays. Al-Nasiri et al. (1975) give a range of 10-13 (9-12, 10 modally but high frequencies at 11 too. Pectoral fin branched rays 16-18 (14-18, modally 16, in Al-Nasiri et al. (1975)), pelvic fin branched rays 8-9, usually 8. Lateral line scales 82-110, lateral line low on the flank anteriorly, rising to the midline of the caudal peduncle. There is a pelvic axillary scale. Scales have a few radii on the posterior field only, a central focus and numerous, fine, concentric circuli. Pharyngeal teeth 3,5-5,3 with variants 2,5-5,3 and 2,5-5,2, long, compressed and hooked at the tip. Gill rakers 9-14, reaching base of adjacent raker when appressed but widely spaced and not developed anteriorly. Some rakers do reach the adjacent one when appressed in some fish. Al-Nasiri et al. (1975) give a range of 11-13 gill rakers with a strong mode at 12. Total vertebrae 51-53 (Al-Nasiri et al. (1975) give 37 as a count which cannot be reconciled with my counts). The gut is an elongate s-shape.

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(4); anal fin branched rays 10(1) or 11(3); pectoral fin branched rays 16(1) or 17(3); pelvic fin branched rays 8(4); lateral line scales 96(1), 98(1) or 100(1); total gill rakers 11(1), 12(2) or 13(1); pharyngeal teeth 3,5-5,3(3); and total vertebrae 51(3) or 53(1).

Sexual dimorphism

Unknown.

Colour

The back is greenish to blackish but overall colour is silvery-grey or silvery-white. Fins are said to be all pale yellow in live fish but are dark in some preserved specimens. A photograph of one freshly caught specimen showed reddish pectoral, pelvic and anal fins, with the dorsal fin greenish, similar to the back and flanks. Another freshly caught specimen was overall silvery, with a brownish-green back, fins overall grey with some yellowish tinges The peritoneum is black to brown.

Size

Reaches over 55.0 cm total length and 6.0 kg in Iraq (van den Eelaart, 1954; Herzog, 1967; Shafi and Jasim, 1982; Bartel et al., 1986) and 1.5 m and 60.0 kg in the Euphrates (Gruvel, 1931; if identification is correct). The Suq al-Shouykh Marsh in April 2005 contained specimens larger than 65.0 cm (www.iraqmarshes.org, downloaded 29 August 2005) and fish in Baghdad palace ponds were estimated to reach 36-40 inches (91-1.02 m) and 15-20 pounds (6.8-9.1 kg) (http://members.cox.net/flybox/FishingUpdate.htm, downloaded 9 January 2006).

Distribution

This species is found in the Tigris-Euphrates and the Orontes River basins in the Middle East. In Iran it is recorded from the lower reaches of rivers in the Tigris River basin including the Bahmanshir River and also such marshes as the Hawr al Azim (Marammazi, 1995).

Zoogeography

This is one of several species that has a sister taxon in the Euro-Mediterranean and/or Black-Caspian-Aral seas basin, indicating north-south connections in the past.

Habitat

van den Eelaart (1954) studied this species in Iraq and found that it lives in rivers, lakes and marshes in both open and vegetated areas and remains in shallow water even in summer. It also occurs in smaller water bodies such as ponds. From spring to fall it is found mainly in marshes and lakes. The barrages at Hindiyah and Kut blocked the upstream migration of this species (Mahdi, 1962). Lakes at Camp Slayer in Baghdad contain this species and, in the shallows, the larger fish chase smaller fish and smaller species leaving v-shaped wakes with the tail fin exposed. Smaller fish leap out of the water to escape the shillik (http://members.cox.net/flybox/FishingUpdate.htm, downloaded 9 January 2006).

Age and growth

Shafi and Jasim (1982) made observations on the biology of this cyprinid in Habbaniyah Reservoir, Iraq. They report 8 age groups with most rapid growth in summer months when water temperatures are above 25°C. Growth in weight is about 160.1 g per year to the fourth year of life and about 331 g per year afterwards. Condition factor was 0.74-1.18 with a mean of 1.0, stable values probably related to piscivory. The length-weight relationship was W = 0.0123 x TL3.0601. The von Bertalanffy equation for growth was lt = 91.0[1-e-0.122 (t-0.25)]. Ali et al. (1986) found the condition factor to range from 0.05 to 1.09 (mean 0.73) and also gave the chemical composition and calorific value. This species had a higher fat content than Barbus (= Carasobarbus) luteus with which it was studied. Al-Dabical and Al-Daham (1995) studied growth in the first year of life in fish from the Shatt al Basrah Canal, Iraq and gave the length-weight relationship as loge W = -12.458 + 3.077 loge L and the growth equation as Lt = 104.118 (1-e -0.0121 (t - 87.871)). Epler et al. (2001) found the oldest age groups to be 5+, 6+ and 7+ in Iraqi lakes Razzazah, Habbaniyah and Tharthar respectively. The mean condition factor was 0.88, 0.76 and 0.87 in lakes Habbaniyah, Tharthar and Razzazah respectively. The von Bertalanffy parameters were for Lake Tharthar L (cm) = 145.5, K = 0.0803, t0 = -0.3269, W (g) = 32099 and n = 3.2249. These indicate rather uniform growth rates, as L is relatively high and K very low. Results were considered more reliable than an earlier study by Jasim (1980) which used inappropriate methods. Annual survival in Lake Tharthar for fish 2.6-5.5 years was 62.0% (Szczerbowski et al., 2001). Productivity was low based on chemical and limnological studies, limiting fish production.

Food

This minnow is piscivorous, feeding almost entirely on fish when adult according to Iraqi studies (Shafi and Jasim, 1982), although aufwuchs may also be found in gut contents. It is mainly a mid-water and benthic feeder with limited predation on surface water organisms (Hussein and Al-Kanaani, 1991). Hussein et al. (1991) examined diet in the Garma Marshes, Iraq and found aquatic insects and crustaceans to be important in young shillig in both summer and winter, with molluscs and fish less important. Even in large shillig, fish were outranked by aquatic insects and in winter by crustaceans as well. Molluscs were a minor food. Shillig rejected certain molluscs while taking others, attributed to variations in shell thickness and a attachment strength to substrates. Liza abu is an important food fish (Al-Shamma'a and Jasim, 1993). Hussein and Al-Kanaani (1989; 1991; 1993) examined the diet of this species in the Al-Hammar Marsh and found a gradually reduced feeding intensity towards the winter months, a highest fullness index in May and lowest in January, and a diet governed by food accessibility and availability. Crustaceans, fish and aquatic insects are the main food items in descending order of importance, with fish most important when using a percentage ranking index in large shillig and even in small shillik by volume. Benthic molluscs were the third most important food for young shillik after crustaceans and fish. In a study of the recovering Hammar Marsh, Iraq, diet was 80.0% fish and 20.0% insects, in the Hawr al Hawizah 47.4% fish and 29.4% insects with shrimps, other crustaceans, algae, diatoms, plants and snails at less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 73.0% fish and 16.8% insects with shrimps, other crustaceans, algae and plants at less than 10% each (Hussain et al., 2006). Fish are the main diet item of large shillik and there is a gradual shift from small- to large-sized prey as the shillik grows (Salman et al., 1994). Frogs, molluscs and aquatic plants and algae were also found in stomach contents, with frogs being important to large shillik in terms of prey volume. Plants may be accidental inclusions taken when seizing prey in weed beds. The fish eaten in descending order of importance were Liza abu, Gambusia affinis (sic, probably G. holbrooki), Garra rufa and Cyprinus carpio. The main crustacean eaten was Metapenaeus affinis along with decapods and amphipods. The gill rakers are widely spaced, indicative of a piscivorous diet (Salman et al., 1993) and the gut is a short s-shape, about equal to fish standard length, also indicative of a piscivorous diet (Salman et al., 1994). Hussain and Ali (2006) examined feeding relationships among fishes in the Al-Hammar Marsh and found this species to be a carnivore, 41.9% of the diet being crustaceans, 10.0% insects and 34.1% fishes. Epler et al. (2001) studied the diet of this species in Lake Tharthar, Iraq and found year old shillik to be eating oligochaetes, tendipedids and plants material with only fish in 2-7 year old shillik. Dietary coincidence with bizz was high in Lake Tharthar, 96.1%.

Reproduction

Shafi and Jasim (1982) record possible spawning in January at 10°C in Iraq with a fecundity up to 74,509 eggs, a mean of 1157 eggs/g body weight and egg diameter of about 1.1 mm. van den Eelaart (1954) found this species in deep parts of Iraqi rivers in December-January, entering marshes and lakes in February to spawn at the end of February and the beginning of March. Spawning takes place on gravel beds, the same as those used by Barbus (= Luciobarbus) xanthopterus, but also on plants. Epler et al. (2001) studied reproduction in Iraqi lakes Tharthar and Habbaniyah and found males to achieve maturity in the third year of life at 44.2 cm and females in the fourth at47.2 cm. Spawning occurred in February at 13-14ºC. Fecundity was 92,000 eggs/kg body mass.

Parasites and predators

Jalali and Molnár (1990a) record the monogeneans Dactylogyrus pulcher and D. mokhayeri from this species in the Dez River. Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 6.6% with the worm Neoechinorhynchus tylosuri in Khuzestan marshes. Farahnak (2000) and Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. It is eaten by Silurus triostegus. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Ergasilus sp., Ergasilus sieboldi, Lernaea sp., Lamproglena sp. and Lamproglena compacta on this species.

Economic importance

Sharma (1980) reports that shillik were an important fish species at the Basrah, Iraq fish market, accounting for 68,948 kg from October 1975 to June 1977, although this is an order of magnitude less than for the three most important species. Its potential for fish farming may be limited by its small gill area which makes it unfit to maintain gas exchange in oxygen-poor water (Salman et al., 1991). Kassim et al. (1998) found locally-raised Scenedesmus acutus algal cultures at 0.5*106 cell/ml with baker's yeast at 0.05 g/L to be the best formula for raising the rotifer Brachionus calcyflorus as live food for shillik larvae. Growth rate was, however, higher on an artificial diet of boiled eggs and soybean meal (52%) compared to 48%, in contrast to common carp (q.v.).

van den Eelaart (1954) gave the fishing season  in Iraq for this species as December-February (peaking in January) and February and June-November (peaking in February and July-August).

Foreign soldiers in Iraq during 2005 regularly caught this species on angling gear using spoons and streamer flies, e.g. www.carpecapio.com, downloaded 26 August 2005.

Conservation

Few specimens have been caught in Iran and deposited in museums. This may reflect rarity or inadequate collection methods. It was commonly caught by American soldiers in Iraq in 2004 as evidenced by emailed photographs sent to me for identification and is an important food fish in Iraq. Detailed surveys using appropriate equipment are needed to assess its distribution and status in Iran. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Its distribution and status in Iran need so be studied as does its distinction from Aspius aspius.

Sources

Scale counts were taken also from Banister (1980).

Type material: See above (NMW 76776 and NMW 76785).

Iranian material: ZMH 2516, 259.9 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh (no further locality data); uncatalogued, 3, 105.6-282.5 mm standard length, Khuzestan, Hawr al Azim and Dez River, (no further locality data). 

Comparative material: NMW 91020, 1, 170.6 mm standard length, Iraq, Shatt-al-Arab, Basrah (30°30'N, 47°47'E); BM(NH) 1920.3.3:127-146, 28, 69.8-284.7 mm standard length, Iraq, Basrah (30°30'N, 47°47'E); BM(NH) 1920.10.8:1, 1, 182.3 mm standard length, Iraq, Tigris River (no other locality data); BM(NH) 1931.12.21:11, 1, 250.2 mm standard length, Iraq, Mosul (36°20'N, 43°08'E); BM(NH) 1972.3.16:1, 1, 112.1 mm standard length, Iraq, Dokan Lake (no other locality data); BM(NH) 1973.5.21:189-190, 2, 166.2-192.0 mm standard length, Iraq, Shatt-al-Arab (no other locality data); FMNH 51242, 1, 322.6 mm standard. length, Iraq, Halfaya east of Amara (31°49'N, 47°26'E); uncatalogued, 1, 200.8 mm standard length, Iraq, Hawr al Hammar (no other locality data). BM(NH) 1968.12.13:182, 1, 251.7 mm standard length, Syria, Cheria River, tributary to the Orontes River (no other locality data); NMW 90366, 1, 309.0 mm standard length, Turkey, Cermik on the Euphrates River (39°09'N, 39°27'E); NMW 90807, 1, 214.8 mm standard length, Turkey, Devegeçidi Çayi, Tigris River basin (no other locality data);

Genus Barbus
Cuvier and Cloquet, 1816

The barbels, genus Barbus sensu lato, are found in Europe, Southwest Asia and Africa and comprise about 800 species with 15 formerly recognised in Iran. Only a single species is now assigned to this genus.

This genus included a wide variety of species and was something of a catchall, serving to cover groups of species which have not been satisfactorily defined as distinct genera to general acceptance. Some authors recognise genera not recognised by others or regard these genera as subgenera - this necessarily affects the species count above. Characters in Southwest Asian species include a rounded or compressed body of moderate to very large size, large to very small scales (lateral line scale count range is at least 26-103), no scale sheath around the anal fin, scales have moderate to high numbers of radii and numerous fine circuli, the presence of barbels in most species, usually 2 pairs, often 1 pair and sometimes none (and individually variable within species), lips variably developed from thin to thick and fleshy, the lower lip sometimes with a well-developed median lobe (and lip development individually variable within species), the last unbranched ray in the short dorsal fin (usually 7-8 branched rays but sometimes more) is thickened and spine-like and may bear teeth or be smooth, a short anal fin, usually with 5 branched rays (but some have 6), pharyngeal teeth in 3 rows with hooked or spoon-shaped tips but sometimes heavy and massive or molariform, gut short, peritoneum white to brown or black, and colour usually brown without distinctive markings in the form of stripes, bands or spots (Luciobarbus subquincunciatus is an exception).

Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003) restrict Barbus to tetraploid species with scales having divergent striae. These species have 7-8, occasionally 9, branched dorsal fin rays, 5 branched anal fin rays, papillose lips and two pairs of barbels. This then excludes species placed in Carasobarbus, Kosswigobarbus, Mesopotamichthys and Tor (see below). Two groups of species can be distinguished in this restricted Barbus according to Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003), namely those with 5 pharyngeal teeth in the main row and a papillose lower lip separated from the chin by a groove and those with 4 pharyngeal teeth in the main row and a lower lip without papillae and continuous with the chin, this latter group being formerly recognised as the genus Luciobarbus Heckel, 1843. The European/Caucasian member(s) of Barbus sensu.stricto in Iran is lacerta and of Luciobarbus (treated as a subgenus in Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003)) are brachycephalus and capito.

Berrebi and Tsigenopoulos in Bănărescu and Bogutskaya (2003) and Tsigenopoulos et al. (2003) review Barbus using molecular markers. They include Barbus cyri (a subspecies of B. lacerta according to some authors) and B. lacerta in the subgenus Barbus, their Northern Mediterranean Group, and B. brachycephalus, capito, esocinus, longiceps, mursa, mystaceus, pectoralis, rajanourum, subquincunciatus, xanthopterus and probably barbulus, kersin, sheich and scincus in the subgenus Luciobarbus, their Southern Group. Levin (2004) studied phenetic relationships of 7 Caucasian taxa and concurred with the division into Barbus and Luciobarbus. See under the species Kosswigobarbus kosswigi for a discussion about the genus/subgenus Kosswigobarbus.

The genus Barbus sensu lato Cuvier and Cloquet, 1816 has been split into a number of genera which are now finding general acceptance. Names used in Southwest Asia include Tor Gray, 1834 sensu Karaman, 1971, Labeobarbus Rüppell, 1836, Systomus McClelland, 1838, Luciobarbus Heckel, 1843, Barynotus Günther, 1868 (preoccupied), Aspiobarbus Berg, 1932, Bertinius Fang, 1943 (and Bertinus Banister, 1980, a misspelling), Bertinichthys Whitley, 1953 (an unneeded replacement of Bertinius), Mesopotamichthys Karaman, 1971, Carasobarbus Karaman, 1971 and Kosswigobarbus Karaman, 1971. Labeobarbus is generally considered to be a synonym of Tor, species of which are found mostly in the Oriental Realm, with only Tor grypus in Iran being a member of the genus Tor (Karaman, 1971; Ekmekçi and Banarescu, 1998). Bertinius is regarded as a synonym of Luciobarbus in Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003). A summary table of generic and/or subgeneric names is given below:-

Species

Original genus

Genus

barbulus

Barbus

Luciobarbus

brachycephalus

Barbus

Luciobarbus

capito

Cyprinus

Luciobarbus

esocinus

Luciobarbus

Luciobarbus

grypus

Barbus

Tor

kersin

Barbus

Luciobarbus

kosswigi

Cyclocheilichthys

Kosswigobarbus

lacerta

Barbus

Barbus

luteus

Systomus

Carasobarbus

mursa

Cyprinus

Luciobarbus

pectoralis

Barbus

Luciobarbus

sharpeyi

Barbus

Mesopotamichthys

sublimus

Barbus

Kosswigobarbus

subquincunciatus

Barbus

Luciobarbus

xanthopterus

Luciobarbus

Luciobarbus

There are also conflicting views on the validity and synonymy of several nominal "Barbus" species. An extensive comparison of these views is not given here (see, for example, Myers (1960), Karaman (1971), Almaça (1983, 1984a, 1984b, 1986, 1990, 1991, 1992, 1994), Krupp (1985c), Howes (1987), Doadrio (1990), Eschmeyer (1990), Berrebi (1995), Berrebi et al. (1996), Tsigenopoulos and Berrebi (2000)). Karaman's studies have not found general acceptance. Author's views conflict, even when examining the same material. Problems include:- the low number of specimens examined (Almaça (1984a; 1986) for example, examined 11 nominal taxa relevant to Iran in detail but averaged only about 6 specimens per taxon, often from a single locality or outside Iranian waters); a wide range in size of individuals of species being compared making age related changes difficult to assess (denticles in the dorsal fin are often lost with age, barbels are shorter, body shape changes, etc); the possibility of sexual dimorphism; possible variation between populations; ecomorphs being recognised as genera (e.g. Luciobarbus was recognised by having 4, as opposed to 5, teeth in the outer pharyngeal tooth row; Bertinius is founded on this condition and development of molar teeth for crushing molluscs - but this may have risen independently in response to an ecological opportunity (see Krupp (1985c)); paedomorphosis and independent origins from a generalised form in different sites (Mina et al., 2001), and the lack of a wide range of new material. An adequate resolution of the systematics of the Barbus sensu lato species in the Tigris-Euphrates basin in particular would require extensive collections of new material from type localities and from the whole basin and comparison of this material with the extant types. Not all types are extant and some that do exist are in poor condition. If this were not complication enough, "Barbus" species are prone to hybridisation with other "Barbus" species and even other genera, further confusing the resolution of the issue. Almaça (1990) cites a hybridization rate of 5.5-6.0% in "Barbus" of the Iberian Peninsula, higher under changed ecological conditions such as the building of dams.

The status of Bertinius longiceps persicus Karaman, 1971 described from the "Karun b. Ahvaz, Persien" (= Karun River at Ahvaz, Khuzestan) on a single specimen is uncertain (lateral line 56-58, gill rakers 22, subterminal mouth, very short barbels, head somewhat higher and suddenly narrowing compared to the type subspecies of the Jordan and Orontes basins, acuminate snout, dorsal fin margin concave). It is not "Barbus" longiceps (F. Krupp, in litt., 1986). The holotype is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H2509).

The roe or eggs of species in these genera have been implicated in poisoning (Halstead, 1967-1970) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death. Sykes (1927) however, in his account of the travels of Sir John Chardin in Persia (first published in 1686) quotes "Barbel.... the Spawn of them especially is dangerous, being a certain and a violent Vomit, by Reason that the Sun never shines on that Fish, and that it breeds in raw Waters; or because they take it with the Nux Vomica or the Vomiting Nut". Najafpour and Coad (2002) report a case of roe poisoning from eggs of Carasobarbus luteus.

Barbels are found in running water of streams and rivers although some may inhabit ponds, springs and lakes. Most show migrations for spawning. A species called soleymani, possibly a "Barbus" species, was considered to be on the verge of extinction in the Gav Masiab River of the Tigris River basin, through pollution, overfishing, dam building, aquaculture, and introduction of exotics (IranMania.com, 29 December 2006). "Barbus" species in Khuzestan are thought to be the intermediate hosts of Heterophyidae flukes found in humans and carnivores (Massoud et al., 1981).

Kazeraani (1994) gives a short account of Iranian "Barbus" species in Farsi. The common names in Farsi for these fishes generally are سس ماهي (= sos, ses or sas mahi, meaning unknown) and زرده پر (= zardehpar), zardek or zardak and ourange or ourenge (in reference to yellow or orange colorations, probably of the fins).

The origin and movements of "palaearctic" or Euro-Mediterranean "Barbus" species in Southwest Asia have been examined by Banarescu (1976; 1977) and Almaça (1984b; 1988; 1990) and these works should be consulted for further details. These works are not cladistic analyses but groupings of species based on morphological similarities and may be subject to criticism on this account.

The origin of the genus "Barbus" according to these authors lies in East Asia and reached the Euro-Mediterranean region by a Siberian route. "Barbus" became extinct in northern East Asia, Siberia and northern Europe when the climate cooled during either the Pliocene or the Quaternary. Europe was colonised during the Oligocene and it is from Europe through Anatolia that Southwest Asia received many of its "palaearctic" "Barbus". This route of entry probably did not occur before the Pliocene because the Syrian-Iranian Sea, the last connection between the Tethys Sea and the Indian Ocean, blocked passage of primary freshwater fishes into what is now Iran and adjacent regions although a connection between a Balkan-Aegean-Anatolian landmass and Iran was possible during the early Miocene (20-17 MYA). A marine transgression 16.8-11.8 MYA flooding the eastern Paratethys and the rise of mountain barriers led to independent evolution of "Barbus" in the Balkan-Aegean-Anatolian landmass and in the Iranian Plateau. During the late Miocene the eastern marine connection of Paratethys closed (11.8-10.5 MYA) allowing an exchange of "Barbus" between Iran and Anatolia, continuous from that time. The Paratethys became an intracontinental sea, the Sarmatian Sea, with a basin encompassing the present Black, Caspian and Aral seas and neighbouring low-lying areas (Bianco, 1990). The Sarmatian Sea freshened as large rivers entered it during the late Miocene and Pliocene, facilitating dispersal of freshwater fishes. A second route of entry for "Barbus" to northern Iran was via southwestern Siberia and the Aral Sea basin during the early to middle Oligocene. Bănărescu and Bogutskaya in Bănărescu and Bogutskaya (2003) agree on an east Asian origin for "Barbus", dispersing across Siberia and western Asia. The group split into two branches, one forming Barbus sensu stricto and using a dispersal route north of the Ponto-Caspian basin and reaching western Europe and another (Luciobarbus) dispersing across the present-day Mediterranean Sea (see above in discussion of Berrebi and Tsigenopoulos in Bănărescu and Bogutskaya (2003) and Tsigenopoulos et al. (2003) for listing of nominal taxa relevant to Iran in these branches or groups).

A recent overview of "Barbus" systematics restricts the genus to Europe, Southwest Asia and Northwest Africa (Berrebi et al., 1996). Barbus sensu stricto is recognised as a lineage which shares morphological characters, has an ancestral tetraploid origin of 2n=100, and has similar karyotypes, biochemical markers and parasites. Genetic studies indicate four groups of species, namely West European and Ponto-Caspian, Iberian, Northwest African and Levantine. Iberian barbels are found in Spain and Portugal and along within the Northwest African barbels share no species with Iran. The West European and Ponto-Caspian barbels include B. brachycephalus, B. capito and B. mursa, and the Levantine barbels include B. barbulus, B. cyri, B. esocinus, B. lacerta, B. pectoralis, B. rajanorum, B. scincus, B. subquincunciatus and B. xanthopterus. The authors make no comments on the validity of these nominal species and only B. brachycephalus has been examined in detail for karyotypes and/or nuclear markers. This work is continuing and the authors advocate various methods. They note that accurate descriptions of many taxa are lacking and that morphology is still the fastest and most cost-efficient way to identify species. Accurate identification is the foundation for all other studies.

Machordom and Doadrio (2001), using ATPase 6 and 8 and cytochrome b, found differentiation in "Barbus" capito and "B". brachycephalus in the Plio-Pleistocene. A clade of the subgenus Luciobarbus was found for species from the Caucasus (as above), Greece and North Africa compared to the Iberian Peninsula, isolation having occurred after the Messinian salinity crisis 5.5 MY ago when the Iberian Peninsula broke away from Africa.

Berrebi et al. (1996) recommend that Barbus-like species which cannot be allocated to a clearly defined genus should be placed in a genus called `Barbus', surrounded by single quotation marks, until the systematic position is elucidated. In the text of Freshwater Fishes of Iran double quotation marks (") are used for accounts that referred to Barbus in the old sense, including all or part of the the species listed here (see table above).

Barbus lacerta
Heckel, 1843

Hevigh River, Caspian Sea basin, August 2011, courtesy of K. Abbasi
Hevigh River, Caspian Sea basin, August 2011, courtesy of K. Abbasi 

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad
Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad
Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad

Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad
Chame Saqez River, Kordestan, Lake Orumiyeh basin, courtesy of A. Mahjoor Azad

Caspian Sea basin, courtesy of A. Mahjoor Azad
Caspian Sea basin, courtesy of A. Mahjoor Azad

Common names

blizem, bellizem, سس ماهي (= sos or sas mahi), زرده پر (= zardehpar), orenge, sos mahi Kura.

[Kur sirbiti in Azerbaijan; murtsa, murza, muruza, muruz in Transcaucasia generally; mursa in Armenia; shabout moraqqat in Arabic in Iraq; karrid or karad achmar (red frill or shag, probably from the colour and the long barbels) and karrid asrak (= blue shaggy one) according to Heckel (1843b) in Arabic in Aleppo; Kurinskii usach or Kura barbel in Russian].

Systematics

Howes (1987) places this species in Barbus sensu stricto. Karaman (1971) assigns many taxa as subspecies of Barbus plebejus Bonaparte, 1832 (dated correctly 1839 in Eschmeyer et al. (1996), see Bianco (1995a) for details), found throughout Europe and Southwest Asia. Bianco (1995a) considers that Barbus plebejus is restricted to Adriatic drainages of Italy and Croatia. Valiallahi (2006) considers B. plebejus to be present in Iran and distinct from B. lacerta based mainly on body shape, the relative head length, the body depth and the fourth dorsal fin ray. Barbus plebejus kosswigi Karaman, 1971 is described as new from the "Oberer Teil des Tigris-Systems" and "Hamam suyu, Beytusebab-Hakkari" (upper Tigris River basin in Turkey). Almaça (1991) considers it an ecophenotype of his Barbus plebejus scincus since two subspecies of the same species cannot live in the same river basin. Barbus plebejus kosswigi is a secondary homonym of Cyclocheilichthys (= Kosswigobarbus) kosswigi according to Kottelat (1997).

Barbus plebejus ciscaucasicus Kessler, 1877 is from the western drainages of the Caspian Sea south to Dagestan but only Barbus plebejus lacerta Heckel, 1843 is found in Iran. It is recognised here as a full species since its relationships to European and other taxa cannot be determined on material available for this study. Bianco and Banarescu (1982) place specimens from the Aras River near Maku, which are probably this species, in Barbus cyclolepis cyri De Filippi, 1865.

Almaça (1981; 1983; 1984a; 1984b, 1986) gives lacerta specific status, distinguishing it from Barbus plebejus by the strong denticulations on the last dorsal fin unbranched ray, lower denticle density, number of scales in transverse rows, shorter head and pectoral fin, longer snout, lower body, the decrease in height of the branched dorsal fin rays is gradual and the profile of the fin is straight, unusual in Barbus with a strongly denticulated dorsal spine. Almaça recognises two subspecies from Iranian drainages:- lacerta from the Tigris-Euphrates basin (and Aleppo) and cyri from the southern Caspian Sea basin. Berg (1948-1949) also refers Caspian Sea basin specimens to Barbus lacerta cyri but in Berg (1949) has cyri from the Tigris River basin too. Saadati (1977) suggests that Lake Orumiyeh basin Barbus lacerta are a distinct subspecies based on higher scale counts there (72-89) than in the Caspian Sea basin. However, B. lacerta as recognised has a wide range in scale counts (see below) and counting methods can differ to include or not supernumerary scales in the lateral line and small scales at the caudal fin base. Fishes resembling B. lacerta from the Namak Lake basin have higher scale counts than Caspian Sea specimens although sample size is too small for a definitive study. Berg (1948-1949) notes that his B. lacerta cyri is subject to extremely wide variations in such characters as body depth, fin and barbel lengths, dorsal spine denticle numbers (even absent in some very large fish) and lateral line scale counts, among others. A large series of specimens would be needed to resolve these problems, allowing for size and sexual variation, new character discoveries and consistent methodologies. Molecular studies might be helpful.

Barbus Lacerta was described from the "Flüssen Kueik bei Aleppo" (Heckel, 1843b).

The following species are synonyms. Barbus Scincus Heckel, 1843 described from "Aleppo" and later from the "Flusse Kueik bei Aleppo" in Heckel (1847a), Barbus cyri De Filippi, 1865 described from the "Kur presso Tiflis" (= Kura River near Tbilisi, Georgia) (including Barbus cyri var. tiflissica Kamenskii, 1899 described from the "Kura bei Tiflis" and Barbus cyri var. chaldanica Kamenskii, 1899 described from the "Andshigan-tschai unweit Chaldan"), Barbus caucasicus Kessler, 1877 from the Kura and Araks rivers and tributaries, Azerbaijan, Barbus toporovanicus Kamenskii, 1899 described from the "Toporavan See" (= Lake Paravani or Taparavani at 41°26'N, 43°48'E, in the upper Kura River basin of Georgia), bortschalinicus Kamenskii, 1899 described from the "schwarze Flüsschen (Das schwarze Flüsschen fällt in die Bortschala, rechter Zufluss des Chram, Nebenfluss der Kura)(tschernaja rjetschka)", Georgia, Barbus sursunicus Kamenskii, 1899 described from "Sursuna in dem Flüsscheu (sic) Kara-tschai, Nebenfluss der Kura, oder ihrem Zuflusse, erbeutet in einer Höhe von ca 3200', zwischen den Seen Tschaldyr-göll und Tuman-göll, dass kleinere aus dem Flüsschen Abastuman-tschai" (Azerbaijan; later in the same article this species is spelt zurzunicus), Barbus armenicus Kamenskii, 1899 described from the "See Tschaldyr-göll, 6522' und den Kars-tschai" (Sildir Gölü and the Kars-chai, Turkey), and Barbus angustatus Kamenskii, 1899 described from the "Kura, bei Borshom". Barbus toporovanicus first appeared in Kamenskii (1887) as a variety of Capoeta fundulus (see Capoeta capoeta). Type localities from Kamenskii (1899) are, obviously, taken from the German text; there is also an accompanying and preceding Russian text with localities in Latin and Russian which are very similar, although in some cases abbreviated.

Heckel (1843), the original describer, recognised Barbus scincus as close to his Barbus lacerta but with a shorter head, sharply decurved forehead, small mouth, and small eyes, all characters not easily quantified without detailed analysis. Berg (1949) placed it in the synonymy of lacerta. Berg's view is followed here; others are described by Almaça (1983; 1984a, 1986) who favours placing scincus as a subspecies of Barbus plebejus as noted above.

The problem with the conclusions above remains, as pointed out earlier, the lack of a wide range of new material and DNA data. Barbus lacerta is most probably a species complex and some of the synonyms above may prove to be distinct, while populations in isolated basins in Iran and elsewhere could be new taxa.

Four syntypes of Barbus lacerta are in the Naturhistorisches Museum Wien (NMW 54227), 1 syntype is in the Senckenberg Museum Frankfurt (SMF 3471, formerly NMW), and 1 syntype is in the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 3236, formerly NMW, 110.3 mm standard length, examined February 2006; F. Krupp, pers. comm., 1985; Eschmeyer et al., 1996; Bogutskaya in Bănărescu and Bogutskaya, 2003). The Vienna card catalogue in 1997 lists one of NMW 54227 as the lectotype. The Vienna catalogue lists 6 specimens. Bogutskaya in Bănărescu and Bogutskaya (2003) designates 54227-1, 181.6 mm standard length, as the lectotype.

Syntypes of Barbus scincus from "Aleppo", the type locality in Heckel (1843b), are reported in the Naturhistorisches Museum Wien by Almaça (1986) and were also examined by me (NMW 22272, 2 specimens, 97.6-146.7 mm standard length, in poor condition and NMW 54526, 1 specimen, 158.8 mm standard length, designated as a lectotype by F. Krupp, 31 October 1984). Eschmeyer et al. (1996) also list NMW 54525 as a syntype and this fish measured 124.2 mm standard length and had been dried at some point before it was examined by me. The Vienna catalogue lists 4 specimens and the card catalogue in 1997 lists these 4 fish with NMW 54526 as "? lectotype" (sic).

Tortonese (1940) and Eschmeyer et al. (1996) list the holotype of Barbus cyri as in the Istituto e Museo di Zoologia della R. Università di Torino (MZUT N.690).

The lectotype of Barbus armenicus, as established by Berg (1948-1949:Fig. 451), is in the Zoological Institute, St. Petersburg under ZISP 5198 with 3 paralectotypes (Eschmeyer et al., 1996).

The lectotype of Barbus sursunicus is in the Zoological Institute, St. Petersburg under ZISP 14740 as established in Berg (1948-1949:fig. 451).

Abdurakhmanov (1962) compares fish from the Aras and Kura river basins and the Lenkoranchai. Lenkoran fish have fewer scales, longer head length and depth, greater maximum body depth, greater anal fin height, longer pelvic and ventral fins, a longer lower caudal fin lobe, a shorter caudal peduncle length, a smaller eye, and a shorter interorbital width than Kura and Aras fish; Lenkoran fish have a longer predorsal distance, greater caudal peduncle depth, and greater dorsal fin height than Kura fish though Aras fish are the same; Lenkoran fish have the dorsal fin base and postorbital distance less than in Aras, but not Kura, fish. No taxonomic distinction is made for these variations.

Key characters

The spotting on the body is characteristic.

Morphology

The mouth is moderate in size, with moderate to thick tuberculate lips. The median lobe of the lower lip is not developed, being small to absent; however the lip does have a central area which is thicker and distinct from the lips laterally in small fish. Bogutskaya in Bănărescu and Bogutskaya (2003) gives illustrations of lower lip development and variations in head shape. Males were thought to have a straight head profile while in females the profile falls steeply in front of the nostrils but Bogutskaya in Bănărescu and Bogutskaya (2003) found some males with a hump on the snout. Morphology is quite variable. Barbels are thick, the anterior one not extending past the nostril level and the posterior one reaching or exceeding the preopercle level.

Dorsal fin with 3-5, usually 4-5, unbranched rays followed by 7-9, usually 8, branched rays, anal fin with 3 unbranched rays followed by 4-6, usually 5, branched rays. Pectoral fin branched rays 13-19 and pelvic fin branched rays 7-8. Lateral line scales 49-87. Scales are a horizontal oval to rectangular in shape with the anterior margin bearing a central protuberance, and sometimes a wavy form. Radii are numerous on all scale fields around a subcentral anterior focus with few to moderate numbers of circuli (as scales are small). Scales may be irregularly arranged on the flank because of their small size giving different counts depending on whether smaller scales are included in the lateral line count. There is a pelvic axillary scale. Gill rakers 5-13, short and just reaching the one adjacent when appressed. Rakers may not develop on the anterior arch giving a wide range in counts. Vertebrae 39-45. Pharyngeal teeth 2,3,5-5,3,2 with variants 2,3,5-5,3,1, 1,3,5-5,3,2, 1,3,5-5,3,1, 2,3,4-5,3,2, 2,3,5-4,4,2, 2,4,5-4,4,4 and even 1,2,3,5-5,3,2,1. The fourth inner row tooth is usually the largest, slightly larger, or slightly smaller in some, than the third. The fifth inner row tooth is blunt and other teeth are hooked or pointed. Teeth may be slightly serrated and there is a short concave surface below the hook. The last unbranched ray of the dorsal fin is moderately to strongly developed, varying between individuals and populations, with denticle density high (up to 65) along three-fifths to two-thirds of its length. Denticle extent appears to be quite variable. Denticles are proportionately larger in small fish. The tip of the last unbranched ray is thin and flexible. Denticles may be absent in large fish. The gut is elongate with about 2 anterior and 1 posterior loops.

Meristics in Iranian fish: dorsal fin branched rays 8(36); anal fin branched rays 5(36); pectoral fin branched rays 14(5), 15(7), 16(16) or 17(8); pelvic fin branched rays 7(8) or 8(28); lateral line scales 53(1), 55(1), 56(3), 59(3), 60(3), 63(3), 64(3), 65(2), 66(1), 67(1), 69(3), 70(1), 72(1), 74(2), 76(2), 79(1), 82(3), 85(1) or 87(1); total gill rakers 6(1), 7(9), 8(10), 9(7), 10(6), 11(1) or 13(1); pharyngeal teeth 2,3,5-5,3,2(18), 1,3,5-5,3,1(1), 2,3,4-5,3,2(1) or 2,3,4-4,3,2(1); and total vertebrae ?.

Sexual dimorphism

Females have shorter barbels than males (Berg, 1948-1949) and females have longer anal and ventral fins (Bogutskaya in Bănărescu and Bogutskaya, 2003). Tubercle development in males caught on 25-26 June consists of minute tubercles thickly developed on the head top, sides and ventrally, lining the margin of anterior belly scales but also 1-2 tubercles in mid-scale, on anterior flank scales numbering 1-4 becoming 1 tubercle on more posterior scales although most mid-flank scales lack tubercles. Lower flank and lower caudal peduncle scales bear a tubercle. Back scales have a unique tuberculation consisting of a line rather than a rounded tubercle. The line lies centrally on the scale and extends from the margin part way along the exposed scale. Behind the dorsal fin the back scales have the central line and one on each side radiating back and up and back and down. Tubercles on the dorsal, caudal and anal fins are small and follow the fin branching. they are weak to absent on the pectoral and pelvic fins but are found on the first unbranched pectoral ray in two rows. Males are a dark gold dorsally and all fins slightly reddish with a gold iridescence when spawning (Bogutskaya in Bănărescu and Bogutskaya, 2003). Spawning females have reddish ventral and anal fins.

Colour

The overall colour is yellowish to olive-grey (possibly bluish according to Heckel (1847a)) with numerous, regular dark-brown to black spots on the back, upper flank and dorsal and caudal fins or irregular mottling. The spots may form a stripe in young fish. In general appearance, fish may be quite light or almost blackish as pigmentation level varies individually. The back is olive-brown to light or reddish-brown and the flanks silvery to yellowish. The belly and lower head surface are white. The iris is dark to silvery with a narrow silver-golden ring. Barbels are white. The dorsal fin bears dark spots and extended lines of dark pigment on the rays and membranes. These are not clearly arranged as bars. The margin of the caudal fin is dark in some fish and there may be a band on mid-fin. The caudal fin is often speckled with dark spots which do not form clear bars. The pectoral fin has dark spots and there are odd dark spots on the pelvic and anal fins. The peritoneum is a light brown with dense but spaced melanophores.

Size

Reaches 37.5 cm and 460 g, possibly to 550 g.

Distribution

This species is found in the Tigris-Euphrates, Quwaiq and Caspian Sea basins as well as some internal basins of Iran. In Iran, it is recorded from the Caspian Sea basin in the Aras River and its tributary the Qareh Su, from the Astara to the Atrak rivers including the Anzali Mordab, the upper Safid River drainage in the Qezel Owzan and Shahrud, in Tajan, Babol, Haraz, Sardab, Aras, Hevigh, Tonekabon, Pol-e Rud and Safid rivers, in the Lake Orumiyeh basin in the middle to upper Talkheh River, Nazlu Chai, Tatavi and Zarrineh rivers, the Tigris River basin, and the Esfahan basin (Dopolan River)(Günther, 1899; Laptev, 1934; Berg, 1949; Holčík and Oláh, 1992; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999; Ghorbani Chafi, 2000; Abdoli, 2000).

Zoogeography

Almaça (1991) considers that this species arose from the first wave of colonisers to enter West Asia from South Europe but is more recent in origin than such Barbus (= Luciobarbus) species as esocinus and xanthopterus originating from southwestern Siberia.

Habitat

This species is found in fresh waters and is not migratory. It avoids muddy bottoms and prefers sandy or stony substrates (Solak, 1977; Bogutskaya in Bănărescu and Bogutskaya, 2003). These habitats are rich in benthos, cool, with rapid currents and well-oxygenated; however it may congregate in slow waters where temperatures reach 26°C.

Age and growth

Solak (1989a) examined a population of this species in the Aras River in Turkey and found up to 5 age groups. Abdurakhmanov (1962) records 5 years as life span in Azerbaijan. Çalişkan et al. (1999) also found 5 age groups in Çıldır Lake, Turkey (for Barbus plebejus, probably this species). Fish in age group 2 dominated and the largest fish attained 320 mm and 550 g. Maturity is attained at 2 years for males and 3 years for females (Bogutskaya in Bănărescu and Bogutskaya, 2003).

In the Tajan River fish showed negative allometric growth with W = 0.0202L2.6787 (Patimar et al., 2012).

Food

Plant remains, crustaceans such as amphipods and insect remains such as chironomids and dragonfly larvae have been found in gut contents. Abdoli (2000) lists Plecoptera, Ephemeroptera and Chironomidae. Algae is also consumed (Bogutskaya in Bănărescu and Bogutskaya, 2003).

Reproduction

Eggs number up to 19,680 and a diameter of 2.3 mm (Abdurakhmanov, 1962; Bogutskaya in Bănărescu and Bogutskaya, 2003). Spawning may occur 2-3 times in a season judging by oocyte sizes in mature ovaries and occurs from the end of April to August, varying with locality, once temperatures reach 14°C, ceasing if the temperature exceeds 20°C (Bogutskaya in Bănărescu and Bogutskaya, 2003). Small Iranian specimens (130.7-157.7 mm standard length) have eggs of 1.0 mm diameter and 1.1 mm on 9 July and 11 May respectively. Larger eggs were noted in a fish caught on 9 July (1.7 mm). The spawning season is probably spring for large fish.

Parasites and predators

Molnár and Jalali (1992) record the monogeneans Dactylogyrus carpathicus and D. linstowi from Barbus plebejus, presumably this species, in the Safid Rud. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from Barbus barbus plebejus, presumably this species. Masoumian et al. (2003) record Myxobolus valdogeli while Pazooki et al. (2003) and Pazooki (2006) record Rhabdochona hellichi, Bothriocephalus gowkongensis, Pseudocapillaria tomentosa, Allocreadium isoporum and Paradiplozoon homoion, all reports from fishes captured in the Tajan and Zarem rivers of Mazandaran. Pazooki et al. (2005) record Trichodina perforata from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus goktschaicus and Gyrodactylus sp. from this fish in Zanjan Province. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Ergasilus sp. and Lernaea sp. on this species. Yakhchali et al. (2011) record the acanthocephalan Neoechinorhynchus sp. from Barbus sp. (presumably B. lacerta) in the Zarrienh River of the Lake Orumiyeh basin, concluding that the fish were not suitable for pond cultivation because of the likelihood of severe economic losses from the parasite.

Economic importance

Not commercially important although it does provide sport in mountain areas of the former U.S.S.R.

Conservation

Kiabi et al. (1999) consider this species to be near threatened in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, medium in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Mostafavi (2007) lists it as near threatened in the Talar River, Mazandaran. Endangered in Turkey (Fricke et al., 2007).

Further work

The various populations of this species require more detailed study, especially with molecular methods, to determine their taxonomy.

Sources

Type material: ?

Iranian material: CMNFI 1970-0559, 9, 39.7-114.3 mm standard length, Azarbayjan- eBakhtari, Baranduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1979-0271, 2, ? mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0289, 1, 131.6 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0449, 2, 85.7-92.2 mm standard length, Azarbayjan-e Khavari, river 18 km from Khalkhal (ca. 37º42'N, ca. 48º27'E); CMNFI 1979-0452, ?, ? mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0468, 7, 30.9-96.1 mm standard length, Mazandaran, Haraz River (36º14'N, 52º22'E); CMNFI 1979-0493, 3, ? mm standard length, Mazandaran, stream in Tajan River drainage (36º19'N, 53º23'E); CMNFI 1979-0557, ?, ? mm standard length, (); CMNFI 1979-0558, ?, ? mm standard length, (); CMNFI 1979-0559, ?, ? mm standard length, (); CMNFI 1979-0785, 2, 115.7-134.8 mm standard length, Aazrabayan-e Bakhtari, Shaher Chay (37º27'N, 34º55'E); CMNFI 1979-0786, 1, 84.1 mm standard length, Azarbayjan-e Khavari, Guru Lake (37º55'N, 46º42'E); CMNFI 1993-0125, 1, 83.1 mm standard length, Kermanshahan, Sarab-e Nilufar (34º24'N, 46º52'E); CMNFI 1993-0126, 2, 157.7 mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 1993-0128, 1, 130.7 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 1993-0136, 1, ?105.5 or 108.2 mm standard length, Mazandaran, Sardabrud (36º39'42"N, 51º22'36"E); CMNFI 2007-0086, 1, 164.4 mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); CMNFI 2007-0087, 2, ? mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0088, 2, ? mm standard length, Azarabyjan-e Khavari, Qareh Su east of Lari (38º30'N, 48º03'E); CMNFI 2007-0093, 1, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0095, 4, 25.9-73.3 mm standard length, Azarbaijan-e Bakhtari, Shahr Chay southwest of Orumiyeh (ca. 37º27'N, ca. 44º56'E); CMNFI 2007-0096, 1, ? mm standard length, Azarbayjan-e Bakhtari, Qasemul River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); CMNFI 2007-0097, 1, ? mm standard length, Azarbayjan-e Bakhtari, Barunduz Chay basin south of Orumiyeh (ca. 37º16'N, ca. 45º08'E); CMNFI 2007-0098, 2, 193.1-227.4 mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0099, 2, 28.9-132.1 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); CMNFI 2007-0100, 1, ? mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0103, 3, 43.6-63.7 mm standard length, Kordestan, Zarineh River basin north of Saqqez (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0104, 2, 54.6-71.2 mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º12'N, ca. 46º18'E); CMNFI 2007-0105, 2, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0106, 1, 99.1 mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 1, 64.6 mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E); CMNFI 2007-0117, 1, ?66.4 mm standard length, Kermanshahan, Gav Masiab basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0117, 1, ?67.2 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0118, 1, ? mm standard length, Kermanshahan, Bid Sorkh River between Sangeh and Kangavar (ca. 34º23'N, ca. 47º52'E); USNM 205931 2, 93.0-115.4 mm standard length, Azarbaijan-e Bakhtari, Baranduz River south of Orumiyeh (37º25'N, 45º05'E); ZMH 2634, 1, 130.5 mm standard length, ?, Haraz River.

Comparative material: BM(NH) 1974.2.22:1236, 1, 113.8 mm standard length, Iraq, Karrid Achmar (no other locality data); BM(NH) 1974.2.22:1327-1328, 2, 121.0-129.9 mm standard length, Iraq (no other locality data); BM(NH) 1974.2.22:1349-1350, 2, 63.1-83.0 mm standard length, Iraq, Qizillja River, Lesser Zab and Serokani near Diana, Rowanduz, Greater Zab (mixed sample); BM(NH) 1974.2.22:1351, 1, 146.8 mm standard length, Iraq, Karrid Asrak (no other locality data).

Genus Barilius
Hamilton, 1822

The members of this genus are found from Pakistan to Thailand with one species in the Tigris-Euphrates and adjacent basins. Their systematics is still poorly understood and there may be about 25 species.

This genus is characterised by a compressed but slender and small body, having small to moderate sized scales, a decurved lateral line, running for example on the lower part of the caudal peduncle, lateral line complete, incomplete or absent, a short dorsal fin and a long anal fin, no fin spines, a moderate and terminal mouth, barbels absent or in 1 or 2 pairs, short gill rakers, pharyngeal teeth in 3 rows, and usually with dark bands or spots on the flank.

These fishes are found mostly in mountain streams although some are lowland species.

Barilius mesopotamicus
Berg, 1932

Common names

None.

[sboura iraqia in Arabic, Mesopotamian minnow].

Systematics

The holotype, 44 mm total length and 35.4 mm standard length, is in the Zoological Institute, St. Petersburg (ZISP 23955) and is decoloured. The collection date is given by Berg (1949) as 16.IV.1914, as 3.IV.1914 in the ZISP catalogue and 5.IV.1914 in the jar. The first two dates are probably correct, one old style and one new style. The type locality is "Stromgebiete des Tigris, in (Siaret) Seid-Hassan, an der persisch-türkischen Grenze, unter 33°20'n. Br., 46°20'ö. L. Seid-Hassan liegt am Flusse Gawi, welcher sich mit dem Kundschian (Gundschian)-tschai vereinigt; der letztere mündet in den Tigris". Seyyed Hasan (33°06'N, 46°11'E) lies on a tributary of the Kanjan Cham River near the Iranian town of Mehran on the Iran-Iraq border. The tributary is presumably the Gawi River.

Howes (1980) stated that this species has apomorph characters shared with species assigned to Leucaspius Heckel and Kner, 1858 but this seems unlikely on general morphological grounds (Coad, 1982b) and Bianco and Banarescu (1982) and Liao et al. (2011) concur, the latter also incorporating molecular evidence. It resembles other Barilius in having barbels (none in Leucaspius), a lateral line low on the body (short and mid-body), broad suborbital bones, and flank bars (none) while Leucaspius is unique in having in females a fold of skin in the shape of two, large, rounded papillae around the genital opening. Bianco and Banarescu (1982) state that this species may be generically distinct from South Asian Barilius but do not diagnose a new genus. Bănărescu and Coad (1991) and Bănărescu (1992b) state that its position and biogeographical affinities are uncertain. Berg (1949) considers it closer to Indian species of the genus Barilius than to African ones.

Key characters

The only member of its genus in Iran, this species is easily identified by the pigment pattern, low lateral line, broad suborbital bones and the barbels.

Morphology

The lower jaw bears a small symphysial knob. The mouth is slightly subterminal, oblique and elongate with the mouth corner under the anterior half of the eye. A well-developed barbel has its origin just anterior to the level of the nostril above the upper lip and lies in a groove between the upper lip and the beginning of the suborbital bone series. This barbel can be absent or minute in some fish (females from Habbaniyah, Iraq (Coad and Krupp, 1983)). In addition to these maxillary barbels, a second pair of barbels have their origin slightly above the posterior edge of the mouth in 8 out of 259 fish examined. They are usually rudimentary but may reach 10.7% of head length. Barbels are difficult to see in smaller fish without magnification. The suborbital bone series is large.

Dorsal fin unbranched rays 2-3, usually 3, branched rays 7-9; anal fin unbranched rays 2-3, usually 3, branched rays 10-14, branched pectoral fin rays 11-15 and branched pelvic fin rays 6-8. Lateral line scales 42-58. Lateral line incomplete or complete, rarely terminating at the pectoral fin level. Lateral line decurved and parallel to the ventral body profile from the pelvic fin origin to the caudal peduncle, being 2-3 scales above this profile. On the caudal peduncle the lateral line is below the mid-line while scales on the caudal fin posterior to the hypural plate are perforated in the mid-line. Pectoral and pelvic axillary scales present. Scales are regularly arranged over the whole body but are not strongly imbricate, particularly on the belly and back anterior to the dorsal fin. Anterior flank scales are oval with subcentral anterior focus and a moderate number of circuli. Radii are found principally on the posterior and lateral fields. Anterior field radii are usually absent although 1-2 radii may occasionally be found. Scale radii based on 5 anterior flank scales from 5 fish (40.7-50.7 mm standard length) number 5-11 primary radii, 0-13 secondary radii and 5-23 total radii. Total gill rakers 7-14. Gill rakers are short and rounded, reaching to or part way to the raker below when appressed. Total vertebrae 38-41. Pharyngeal teeth usually 4,5-5,4, often 4,5-5,3 (25% of 20 fish examined), or more rarely in three rows 1,3,5-5,3,1 or 1,4,5-5,4,1. Teeth are hooked at the tip, slender and have a concave grinding surface below the tip. The gut is a simple s-shape.

Meristic values for Iranian specimens are:- branched dorsal fin rays 7(2), 8(32) or 9(2); branched anal fin rays 10(2), 11(19), 12(13) 13(1) or 14(1); branched pectoral rays 11(1), 12(8), 13(23), 14(2), or 15(1); pelvic fin rays 6(1), 7(33) or 8(2); scales in lateral series 42(1), 43(3), 44(3), 45(2), 46(6), 47(6), 48(1), 49(3), 50(4), 51(4), 52(2) or 54(1); total gill rakers 7(4), 8(6), 9(6), 10(7), 11(6), 12(1), 13(1) or 14(1); pharyngeal teeth 4,5-5,4(8), 4,5-5,3(3) or 1,3,5-5,3,1(1); and total vertebrae 39(8), 40(20) or 41(5).

Sexual dimorphism

Unknown.

Colour

Overall colour is a brilliant silver with a golden-yellow glimmer, with the back darker and having a thin median stripe. Scales are highly deciduous and leave a silvery smear on the hand. The flanks have 6-11 roundish dark, grey-green spots, not clearly apparent in live fish. In preserved fish the spots are brown. A median dorsal stripe is variably developed. Fins are lightly pigmented, most melanophores being on the rays rather than the membranes. The anal and paired fins are almost entirely hyaline. The caudal fin may show one or two irregular bars running parallel to the posterior margin. The peritoneum is light to silvery but bears scattered melanophores which give a greyish tinge in preserved fish. Some fish from Iraq (Habbaniyah stream) lacked, or had weakly expressed, flank spots.

Size

Reaches 50.7 mm standard length.

Distribution

This species is found in the Tigris-Euphrates basin, including its Iranian part and the adjacent Gulf basin (Berg, 1932; 1949; Bianco and Banarescu, 1982). Abdoli (2000) also records this species from the Jarrahi, the lower Karun, the lower Dez, the Zohreh, the lower half of the Helleh, and the middle and lower Mand rivers.

Zoogeography

This species is found in the Tigris-Euphrates basin of Turkey, Syria, Iraq and Iran. It does not appear to be common in Turkey, at least in the upper reaches of this basin there, nor in upper reaches of Iranian rivers. The distribution in the Dalaki River of Iran is outside the modern Tigris-Euphrates basin. It is presumably a relict of the late Pleistocene when the Tigris-Euphrates flowed down a drained Gulf receiving tributaries now isolated by the post-Pleistocene rise in sea level (Coad and Krupp, 1983).

Habitat

Found in both running and still water, from small streams only 1 m wide and irrigation ditches to major rivers more than 200 m across. Current is slow to fast but generally an obvious flow is apparent. However one specimen was collected in a fish pond near Ahvaz (ZSM 25701). The collection localities in Iran are all at low altitudes and no fish were taken in Zagros Mountain streams and rivers. Collections were made over mud and pebble substrates in shallow streams or at river margins. The species may also occur at the surface in mid-river but no collections confirm this supposition. Capture temperatures were 12-24°C and conductivity 0.45-10.5 mS. Salinity in drying pools of 20 cm depth in Syria where this species was caught in March had Cl-1 = 390 mg/l and a salinity of 1.5‰ (Coad and Krupp, 1983).

Age and growth

Unknown.

Food

Gut contents include winged insects (Coleoptera, Heteroptera, Thysanoptera and Diptera) and spiders, suggestive of surface feeding (Coad and Krupp, 1983). Abdoli (2000) also reports Hymenoptera, Brachycera and Culicidae.

Reproduction

Most fish were collected in January when eggs were small but developing suggestive of spring spawning. Al-Rudainy (2008) gives an absolute fecundity of about 200 eggs for Iraq.

aParasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This fish is found in suitable habitats of large rivers and in small ditches and does not appear to be in need of conservation. Vulnerable in Turkey (Fricke et al., 2007).

Further work

Molecular or detailed osteological analyses might reveal its relationships to taxa from the Oriental region.

Sources

Type material: See above (ZISP 23955).

Iranian material: CMNFI, 1979-0120, 3, 19.3-50.7 mm standard length, Bushehr, Dalaki River near Konar Takhteh (29º28'N, 51º21'E); CMNFI 1979-0357, 1, 27.6 mm standard length, Khuzestan, Karkheh River drainage (31º34'N, 48º12'E); CMNFI 1979-0363, 11, 21.4-30.2 mm standard length, Khuzestan, Karkheh River (31º52'N, 48º20'E); CMNFI 1979-0365, 7, 20.0-34.4 mm standard length, Khuzestan, Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0367, 1, 34.2 mm standard length, Khuzestan, Meymeh River (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 29, 21.6-41.9 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0372, 2, 30.7-33.1 mm standard length, Khuzestan, Dez River near Chogha Zanbil (ca. 32º02'N, ca. 48º30'E); CMNFI 1979-0377, 3, 28.0-39.4 mm standard length, Khuzestan, Karkheh River (ca. 32º57'N, ca. 47º50'E); CMNFI 1979-0378, 7, 31.9-42.4 mm standard length, Khuzestan, stream tributary to Karkheh River (ca.32º48'N, ca. 48º04'E); CMNFI 1979-0380, 10, 25.3-41.0 mm standard length, Khuzestan, stream tributary to Dez River (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0381, 7, 24.3-31.2 mm standard length, Khuzestan, stream west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0382, 4, 25.9-30.8 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 8, 28.6-34.8 mm standard length, Khuzestan, Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 3, 26.8-40.8 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0392, 3, 35.0-39.3 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0396, 35, 25.1-48.8 mm standard length, Khuzestan, Kheyrabad River (30º32'N, 50º23'30"E); ZSM 25701, 1, 36.5 mm standard length, Khuzestan, fishpond near Ahvaz (no other locality data); ISSB uncatalogued, 1, 48.7 mm standard length, Bushehr, Helleh River (ca. 29º20'N, ca. 51º15'E) (Coad and Krupp, 1983).

Comparative material:- BM(NH) 1974.2.22:1256-1267, 11, 33.7-46.2 mm standard length, Iraq, stream between Lake Habanniyah and Euphrates River (ca. 33º22'N, 43º34'E); BM(NH) 1968.12.13:217-220, 4, 18.5-47.4 mm standard length, Syria, Euphrates River at Mayadine (35º01'N, 40º27'E); BM(NH) 1968.12.13:221-236, 16, 30.8-42.4 mm standard length, Syria, Tigris River at Ain Diwar (37º17'N, 42º11'E); SMF 16442, 5, 28.2-35.9 mm standard length, Syria, Nahr Balikh at Jisr Shanine (36º03'N, 39º06'E); SMF 16443, 63, 17.0-34.9 mm standard length, Syria, Nahr Balikh at Jisr Shanine (36º03'N, 39º06'E); ISSB uncatalogued, 4, 32.8-34.4 mm standard length, Turkey, Batman Suyu (ca. 37º55'N, ca. 40º15'E) (Coad and Krupp, 1983).

Genus Blicca
Heckel, 1843

Shutov (1969) places this genus and species in the genus Abramis Cuvier, 1817 on the basis of literature data as does analyses by Shcherbukha (1973) and Howes (1981). Hensel (1978) and Tadajewska (1998) also place this genus in Abramis on the basis of the lateral line system structure, pharyngeal teeth, scale and dermal bone morphology along with data on ecology, behaviour, ontogenesis, osteology and parasitofauna. Hänfling and Brandl (2000) consider Blicca a junior synonym to Abramis based on allozyme data. In contrast, Bogutskaya (1986) using skull morphology reaffirms its generic status.

The white bream genus contains a single species found from Europe to the Caspian Sea basin including Iran.

The genus is characterised by a deep and strongly compressed body; scales absent on the back behind the dorsal fin thus forming a narrow groove; a scaleless keel between the vent and the pelvic fins; pharyngeal teeth in 2 rows; a small, oblique and subterminal mouth; moderate number of gill rakers; scales of moderate size; a short and spineless dorsal fin and a long anal fin; and a light peritoneum.

Blicca bjoerkna
(Linnaeus, 1758)

Anzali Wetland, November 2011, courtesy of K. Abbasi
Anzali Wetland, November 2011, courtesy of K. Abbasi

Blicca bjoerkna, Wikimedia Commons
Blicca bjoerkna, Wikimedia Commons

Common names

simparak or seamparak (= silver scales, possible meaning since parak is a small feather), سيم نما (sim nama or mahi sim nama, meaning silvery-like fish or sim-like fish in reference to Abramis brama).

[yastigarin in Azerbaijan; Zakavkazskaya gustera or Transcaucasian white bream, Armyanskaya gustera for A. b. derjavini, all in Russian; silver bream, white bream, flat bream].

Systematics

Cyprinus Björkna was originally described from Lake Mälar, Sweden.

Cyprinus Blicca Bloch, 1782 described from lakes in Germany, Cyprinus gibbosus Pallas, 1814 described from the Sura and Volga rivers and Blicca argyroleuca Heckel, 1843 are synonyms. It appears that the latter taxon is first described in Heckel's work on fishes of Syria, but in the section devoted to classification based on the pharyngeal teeth of cyprinids; the taxon is later described from Europe in Heckel and Kner (1858) and is not a Southwest Asian species. Syntypes of Blicca argyroleuca are in the Naturhistorisches Museum Wien under NMW 16901 (2 fish), NMW 54918 (6), NMW 54919 (4) and NMW 54920 (1) (Eschmeyer et al., 1996). The spelling bjorkna is incorrect (Eschmeyer et al., 1996).

The Caspian Sea basin subspecies is Blicca bjoerkna transcaucasica Berg, 1916, described from the lower reaches of the Kura River, Araks, Lenkoran District, Transcaucasia. It is distinguished by "somewhat" fewer rays in the anal fin (17-21) and "a tendency to have" fewer lateral line scales (40-45) than in the type form which mostly has 21-22 anal fin rays and 45-48 lateral line scales (Berg, 1948-1949). Abdurakhmanov (1962) expands these ranges to 17-22 and 40-48 respectively but gives low means (± standard error) for 100 fish from Azerbaijan of 19.88±0.13 and 43.56±0.05 respectively. This may be a valid subspecies but the possibility of clinal variation has not been examined.

Blicca bjoerkna derjavini Dadikyan, 1970 is described from the "Sevdzhur River, (tributary of Araks River, in Armenian SSR) and the canal and lake system connected with it". It is distinguished from transcaucasica by lower mean number of branched dorsal fin rays and branched anal rays, a higher mean lateral line scale count, and various morphometric characters.

Key characters

The scaleless ventral keel, postdorsal groove, long anal fin, lateral line scale count and small and oblique mouth are characteristic.

Morphology

Dorsal fin with 3 unbranched and 7-10 branched rays, usually 8, anal fin with 3 unbranched and 16-24 branched rays. Pectoral fin branched rays 14-16, pelvic fin branched rays 7-9. Lateral line scales 40-55. Scales have numerous fine circuli, an almost central focus, a wavy anterior margin and a crenulate posterior margin, and few primary anterior and posterior radii, as few as 2 in each field (there may be numerous secondary radii which do not reach the focus). There is a pelvic axillary scale. Gill rakers 12-21, touching the adjacent raker when appressed. Vertebrae 37-43. Pharyngeal teeth 2,5-5,2 with variants 2,5-5,1, 1,5-5,2, 1,5-5,1, 2,5-4,2, 2,5-4,1, 1,5-4,1, 3,5-5,2, and 3,5-5,3 (among others, see below and Tadajewska (1998)), weakly hooked (strongly hooked in young), compressed, concave below the tip and smooth (anterior tooth margin serrated in young). In young fish, the first major row tooth may be medial to the second tooth rather than in line. Tadajewska (1998) gives details of tooth development. The intestine is s-shaped with a small anterior loop. The chromosome number is 2n=50 (Klinkhardt et al., 1995; Pourkazemi et al., 2010).

Meristic values for Iranian specimens are:- dorsal fin branched rays 8(49) or 9(1); anal fin branched rays 17(3), 18(15), 19(21), 20(9) or 21(2); pectoral fin branched rays 14(15), 15(24) or 16(7); pelvic fin branched rays 7(1), 8(47) or 9(2); lateral line scales 41(2), 42(10), 43(9), 44(13), 45(10), 46(5) or 47(1); total gill rakers 13(2), 14(25), 15(18), 16(4) or 18(1); pharyngeal teeth 2,5-5,2(3), 2,5-5,1(4), 1,5-5,2(4), 2,5-4,1(1), 2,5-5,0(1), 0,5-5,2(1), 1,5-5,1(1), 1,5-5,0(2), 1,5-4,1(2) or 2,4-4,1(1); and total vertebrae 38(8), 39(33) or 40(12).

Sexual dimorphism

Breeding males have fine tubercles on the top of the head, operculum and lining the exposed scale margins on the flank. There are occasionally tubercles in mid-scale. Small tubercles are found on the pectoral fin rays, 1-3 rows on the unbranched ray, 1-2 on the first branched ray and usually 1 on the other rays, branching to follow the branching rays. Other fins bear fine tubercles following the fin rays. Larger tubercles are found in clumps on the scales overlapping the anal fin base. Tubercles are absent from the belly. Fine unculi are present on the snout, under the eye and between the tubercles on the head generally as well as on the underside of the pectoral fin.

Colour

The back is a bluish-green and the rest of the body silvery. The pectoral and pelvic fins are orange-red with grey tips. The peritoneum is silvery with scattered melanophores.

Size

Reaches 54.5 cm and 2.13 kg, possibly 2.25 kg (Machacek (1983-2012), downloaded 27 July 2012).

Distribution

Found from England through Europe north of the Alps and Pyrenees to the Caspian Sea basin. Apparently it does not penetrate to the higher reaches of even major rivers like the Kura and Aras. In Iran it is found from the Aras River (including its middle reaches in Iran) to the Atrak River in the Caspian Sea basin including the Gorgan, Tajan, Babol, Haraz, Sardab, Tonekabon, and Safid rivers, the Anzali Talab and the Boojagh Wetland (Derzhavin, 1934; Holčík and Oláh, 1992; Nejatsanatee, 1994; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009; Khara et al., 2011).

Zoogeography

This species is part of a northern European and northern Southwest Asian fauna whose zoogeographical history has not been thoroughly researched. The relationships with similar genera are reviewed under the genus.

Habitat

This species is found in the shallows of warm lakes with heavy vegetation and in the slower reaches of rivers including river estuaries in Iran (Jolodar and Abdoli, 2004). It overwinters in deeper water. There was a mass mortality of this species on the Babol Sar beach on 24 June 1963 (USNM 271217).

Age and growth

Growth is slow with maturity attained at 3-5 years and 10-12 cm. Some males may mature at 2 years. Females are much larger than males of the same age. Life span is up to 16 years. Stunted populations comprising large numbers of individuals develop where predators are absent.

Food

Food items include insect larvae such as chironomids, worms and molluscs, and some vegetation. This is a euryphagous species. Young fish feed principally on copepods and cladocerans. Even adults will feed on plankton and it is less of a bottom feeder than Abramis brama.

Reproduction

Spawning in the Volga delta takes place about the beginning of May at around 11°C water temperature but may run from the end of April to the middle of July in the Volga generally. Spawning in the Aras flood plain occurs in the middle of April. Generally spawning occurs later than in Abramis brama and Rutilus rutilus but may overlap and infertile hybrids result. Shallow weedy areas are preferred. Each female is pursued by several males. Fecundity reaches 109,000 eggs and egg diameter 1.44 mm. Eggs adhere to plants or stones on the bottom. There can be 3 spawnings at intervals of 10-11 days when water temperatures are at least 16-17°C. Batch spawning shows much individual variation as well as varying between localities and by year at the same locality.

Parasites and predators

Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Tajbakhsh et al. (2010) report the nemtode Philometra rischta from fish in the Anzali wetland. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species. Khara et al. (2011) list the mongenean Dactylogyrus sp. and the crustacean Lernaea cyprinacea from this fish in the Boojagh Wetland of the Caspian Sea.

The Caspian seal, Pusa caspica, is a predator on this species (Krylov, 1984) as are a variety of other fishes such as perch (Perca fluviatilis) and pike-perch (Sander sp.). Ashoori et al. (2012) found that grey herons (Ardea cinerea) in the Siahkeshim Protected Area of the Anzali Wetland ate this species.

Economic importance

Holčík and Oláh (1992) report a catch of 144 kg in the Anzali Mordab in 1990.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and in textbooks.

Conservation

Lelek (1987) classifies this species as intermediate in Europe (liable to be transferred to vulnerable or rare categories if their habitat deteriorates further). Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), absent in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The biology of this species needs study in Iran.

Sources

CMNFI 1970-0510, 1, 56.0 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0522, 4, 40.0-62.6 mm standard length, Gilan, Safid River at Astaneh Bridge (36º16'30"N, 49º56'E); CMNFI 1970-0532, 6, 30.0-63.2 mm standard length, Gilan, Caspian Sea near Bandar-e Anzali (37º28'N, 49º27'E); CMNFI 1970-0553, 4, 62.1-80.1 mm standard length, Gilan, Sowsar Roga River (37º27'N, 49º30'E); CMNFI 1970-0579, 2, 52.6-56.9 mm standard length, Gilan, Old Safid River estuary (37º23'N, 50º11'E); CMNFI 1970-0580, 31, 31.8-86.3 mm standard length, Mazandaran, river near Iz Deh (36º36'N, 52º07'E); CMNFI 1970-0582, 1, 70.9 mm standard length, Mazandaran, Aliabad Reservoir (36º56'N, 54º50'E); CMNFI 1970-0585, 39, 32.4-52.5 mm standard length, Gilan, Nahang Roga River (37º28'N, 49º28'E); CMNFI 1970-0587, 36, 34.6-55.4 mm standard length, Mazandaran, Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0470, 2, 44.5-51.2 mm standard length, Mazandaran, stream west of Alamdeh (36º35'N, 51º43'E); CMNFI 1979-0472, 30, 38.7-69.6 mm standard length, Mazandaran, stream west of Mahmudabad (36º37'N, 52º12'E); CMNFI 1979-0685, 3, 63.1-67.1 mm standard length, Gilan, Safid River (37º24'N, 49º58'E): CMNFI 1980-0117, 1, 80.0 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1980-0122, 15, 38.7-45.3 mm standard length, Mazandaran, Nerissi River (36º38'N, 52º16'E); CMNFI 1980-0149, 6, 60.1-63.7 mm standard length, Gilan, Chabak River (37º21'N, 49º50'E).

Genus Capoeta
Valenciennes, 1842

The genus Capoeta has a wide distribution in Southwest Asia and contains about 20 species of which 7 occur in Iran. Its affinities are uncertain and may lie with the European Barbus/Aulopyge group or with Cyprinion and its southern and east Asian relatives (Karaman, 1971; Howes, 1982; Krupp, 1985c; Bănărescu, 1992b).

Varicorhinus Rüppell, 1836 (as used for Southwest Asian cyprinids) is a synonym of Capoeta Valenciennes, 1842 (see Karaman (1969) for further details: Capoeta is distinguished from Varicorhinus of Africa since it has a denticulate last unbranched dorsal fin ray (as opposed to smooth), very small to medium-sized scales (large), lachrymal bone narrow and covering only a small part of the upper side of the rostrum (large and covering most of the rostrum), suborbital bones narrow and long (short and wide), posterior maxillary process not extending back to a level with the centre of the jugal (extends back to a level of the centre of the suborbitals), lower jaw long (short). Scaphiodon Heckel, 1843 has been used for Capoeta and Cyprinion species in Southwest Asia. The nomenclatural status of this genus is reviewed by Bănărescu in Bănărescu (1999).

Levin et al. (2012) using cytochrome b found that Capoeta is a monophyletic clade nested within Luciobarbus, with origins in the Middle Miocene of a palaeo-Tigris-Euphrates basin. Luciobarbus subquincunciatus is the closest relative and is only found in the modern Tigris-Euphrates basin. The specialised algae scraping morphology appeared once within in the evolution of this genus. Three main groups were detected, the Mesopotamian group (barroisi and trutta in Iran), the Anatolian-Iranian group (buhsei, damascina, saadii), and and the Aralo-Caspian group (aculeata, heratensis). The origin of the genus was most probably through allopolyploidisation, as species are hexaploids (2n = 150). The separation of the Mesopotamian clade occurred in the Middle Miocene about 12.6 MYA, of the Iranian members of the Anatolian-Iranian group in the Pliocene soon after 6.7 MYA, and of the Aralo-Caspian group during the later Pliocene 2.6 MYA.

This genus Capoeta is characterised by a compressed to rounded and moderately elongate body, small to moderately large scales (lateral line counts 37-99), scales at the anal fin base and anus not usually enlarged (sometimes variably enlarged as is the case with certain cyprinids), an inferior, transverse mouth, the lower jaw with a sharp, horny sheath, barbels absent or in 1 or 2 pairs, dorsal fin short (usually 7-9 branched rays) with the last unbranched ray thickened and bearing serrations (serrations sometimes reduced to absent), anal fin short (usually 5 branched rays), gill rakers short, moderate to numerous, pharyngeal teeth in 3 rows with spoon-shaped and truncate tips, a very long and coiled gut (ca. 7-10 times body length), mostly of uniform colour, and a black peritoneum.

The general name for the members of this genus in northern Iran is سياه ماهي (= siah mahi, meaning black fish) while in the south they are called twiny or touyeni and even gel cheragh (= mud-eater, mud-grazer). The name Capoeta is derived from the Armenian and Georgian name for female Capoeta capoeta packed with eggs, namely "Kapwaeti". Other general names for members of this genus shol khar, ghel khar or choul khar, all variant spoken intonations meaning mud eater.

The origin of Capoeta in Southwest Asia follows the same route as the genus Barbus (q.v.).

CMNFI 1977-0510A, 4, mm standard length, Fars, qanat at Naqsh-e Rostam (29º59'30"N, 52º54'E); CMNFI 1979-0026, , mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0027, , mm standard length, Fars, Chehel Chashmeh (ca. 29º43'N, ca. 52º02'E); CMNFI 1979-0036, 2, 83.9-118.3 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0043, , mm standard length, Fars, qanat behind Sarvestan (29º16'N, 53º14'E); CMNFI 1979-0044, , mm standard length, Fars, qanat at Mian Jangal (29º09'N, 53º27'E); CMNFI 1979-0053, 6, 47.3-79.5 mm standard length, Fars, Shur River tributary (ca. 28-29º58-03'N, ca. 52º34-35'E); CMNFI 1979-0054, 16, 35.8-127.9 mm standard length, Fars, Shur River tributary (ca. 28-29º58-03'N, ca. 52º34-35'E); CMNFI 1979-0057, , mm standard length, Fars, Shapur River 4 km from Shapur (29º49'N, 51º34'E); CMNFI 1979-0058, 6, 75.6-115.3 mm standard length, Fars, jube over Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0059, 2, 45.0-50.4 mm standard length, ID, more than one species? Fars, Pulver River 8km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, , mm standard length, ID, more than one species? Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0063, 2, 201.0-206.7 mm standard length, Fars, qanat under Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 1979-0067, , mm standard length, Fars, qanat at Zarqan (ca. 29º46'N, ca. 52º43'E); CMNFI 1979-0068, , mm standard length, Fars, qanat 12 km from Shiraz on Esfahan road (ca. 29º43'N, ca. 52º34'30"E); CMNFI 1979-0073, 5, 28.9-86.6 mm standard length, Fars, Mand River beyond Chehel Chashhmeh (ca. 29º42'30"N, ca. 52º01'30"E); CMNFI 1979-0074, , mm standard length, Fars, Mand River backwater (29º41'N, 52º06'E); CMNFI 1979-0075, , mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0079, 2, 120.7-149.9 mm standard length, Fars, Mand River 5 km above Band-e Bahman (ca. 29º12'N, ca. 52º38'E); CMNFI 1979-0079, 1, 159.7 mm standard length, Fars, Mand River 5 km above Band-e Bahman (ca. 29º12'N, ca. 52º38'E); CMNFI 1979-0090, , mm standard length, Esfahan, Gav Khuni (ca. 32º21'N, ca. 52º49'E); CMNFI 1979-0093, 1, 73.9 mm standard length, (); BWC 76-77 check CMNFI # CMNFI 1979-0109, 1, 91.1 mm standard length, Fars, Mand River ar Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0109, 1, 103.4 mm standard length, Fars, Mand River ar Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0111, 10, 8.7-54.6 mm standard length, Fars, stream 21-22 km from Shiraz (29º37'30"N, 52º21'E); CMNFI 1979-0113, , mm standard length, Fars, qanat under Sa'adi's Tomb (29º37'N, 52º35'E); CMNFI 1979-0114, , mm standard length, Fars, Mand River at road bridge (29º41'N, 52º06'E); CMNFI 1979-0115, 4, 154.4-172.6 mm standard length, qanat under Sa'adi's Tomb (29º37'N, 52º35'E); CMNFI 1979-0125, 1, 137.8 mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0128, 16, 34.6-108.6 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0128, 18, 17.2-135.3 mm standard length, Fars, Shur River (28º51'N, 52º31'E); CMNFI 1979-0129, , mm standard length, Fars, spring 2 km north of Farrashband (28º54'N, 52º04'E); CMNFI 1979-0130, 5, 44.4-93.3 mm standard length, Fars, Shur River 4 km west of Firuzabad (28º51'N, 52º32'E); CMNFI 1979-0131, 58, 25.5-140.0 mm standard length, Fars, Mand River tributary (28º38'N, 52º49'E); CMNFI 1979-0132, 23, 51.1-74.4 mm standard length, Fars, Mand River tributary (28º35'N, 52º58'E); CMNFI 1979-0154B, 6, mm standard length, Fars, upper Shur River drainage near Darab (28º45'30"N, 52º24'E); CMNFI 1979-0155, 7, 36.2-80.5 mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 3, 54.6-122.9 mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, , mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0158, , mm standard length, Fars, qanat over Qasook River (28º54'N, 53º53'30"E); CMNFI 1979-0159, 87, 23.1-167.3 mm standard length, Fars, qanat at Qaziabad (ca. 28º54'N, ca. 53º43'E); CMNFI 1979-0160, 4, 66.3-138.4 mm standard length, Fars, Arteshkadeh Pomp spring (29º09'N, 53º37'E); CMNFI 1979-0161, 29, 33.2-88.3 mm standard length, Fars, qanat on Neyriz to Shiraz road (29º10'30"N, 53º41'E); CMNFI 1979-0162, 9, ?-88.3 mm standard length, Fars, qanat behind Sarvestan (29º16'30"N, 53º14'E); CMNFI 1979-0163, 1, 73.8 mm standard length, ?Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0164, 1, 49.4 mm standard length, ?Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0165, 7, 30.0-96.6 mm standard length, Kerman, qanat at Ahmadabad (30º32'N, 55º38'E); CMNFI 1979-0166, 67, 37.1-123.1 mm standard length, Kerman, qanat at Hassanabad-e Nuq (30º43'N, 55º50'E); CMNFI 1979-0168, , mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0169, , mm standard length, Kerman, qanat 10 km from Mahan (30º08'30"N, 57º17'E); CMNFI 1979-0170, , mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 1979-0171, , mm standard length, Kerman, qanat at Bardesir (29º56'N, 56º34'E); CMNFI 1979-0187, , mm standard length, Hormozgan, stream and pools at Sar Khun (27º23'30"N, 56º26'E); CMNFI 1979-0191, , mm standard length, Fars, stream 10 km east of Furg (ca. 28º16'N, ca. 55º18'E); CMNFI 1979-0192, , mm standard length, Fars, qanat 2 km east of Rostaq (28º26'30"N, 55º04'E); CMNFI 1979-0195, , mm standard length, Fars, jube on road to Fasa (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0198, , mm standard length, Fars, stream at Tadovan (28º47'N, 53º24'30"E); CMNFI 1979-0199, 6, 70.8-102.1 mm standard length, Fars, qanat 18 km from Jahrom (ca. 28º23-25'N, ca. 53º31-40'E); CMNFI 1979-0202, , mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0203, , mm standard length, Fars, qanat at Dudej (29º33'N, 52º59'E); CMNFI 1979-0204, , mm standard length, Fars, qanat on road to Kharameh (29º33'N, 52º59'E); CMNFI 1979-0205, 12, 45.9-200.5 mm standard length, Fars, jube at Runiz-e Pa'in (29º12'N, 53º42'E); CMNFI 1979-0206, , mm standard length, Fars, qanat on road to Kharameh (29º12'N, 53º40'E); CMNFI 1979-0207, 12, 24.2-83.7 mm standard length, Fars, jube 22 km from Neyriz (29º16'N, 54º28'E); CMNFI 1979-0208, 6, 39.9-130.4 mm standard length, Fars, qanat 47 km from Neyriz (ca. 29º11'N, ca. 54º40'E); CMNFI 1979-0209, 60, 43.6-138.9 mm standard length, Kerman, qanat at Kuch Kuluh (29º25'N, 56º03'E); CMNFI 1979-0211, 63, 33.2-94.3 mm standard length, Kerman, river on road to Baft (29º19'N, 56º12'E); CMNFI 1979-0212, 73, 26.0-99.1 mm standard length, Kerman, qanat on road to Baft (29º14'N, 56º17'E); CMNFI 1979-0213, 5, 51.4-60.2 mm standard length, Kerman, stream in Kharan River drainage (29º15'N, 56º25'E); CMNFI 1979-0214, , mm standard length, Kerman, qanat pool on road to Baft (ca. 29º15'N, ca. 56º28'E); CMNFI 1979-0215, 15, 39.7-125.9 mm standard length, Kerman, Kharan River drainage (29º14'N, 56º37'E); CMNFI 1979-0216, 11?, 51.1-65.8 mm standard length, Kerman, qanat 9 km from Baft (ca. 29º13'N, ca. 56º42'E); CMNFI 1979-0217, 15, 39.7-125.9 mm standard length, Kerman, Kharan River drainage (ca. 28º59'30"N, ca. 56º51'30"E); CMNFI 1979-0221, , mm standard length, Kerman, Halil River drainage (28º51'N, 57º52'E); CMNFI 1979-0241, , mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0243, , mm standard length, Esfahan, Zayandeh River at Falavarjan (32º33'N, 51º31'E); CMNFI 1979-0246, , mm standard length, Shahrestan-e Bakhtiari va Chahar Mahall, upper Karun River drainage (31º57'30"N, 50º59'E); CMNFI 1979-0251, , mm standard length, Esfahan, stream 1 km east of Daran (32º59'N, 50º26'E); CMNFI 1979-0251, , mm standard length, Esfahan, stream 1 km east of Daran (32º59'N, 50º26'E); CMNFI 1979-0255, , mm standard length, Markazi, Bar River drainage 2 km west of Shahabiyeh (33º51'30"N, 50º23'E); CMNFI 1979-0269, 1, 125.0 mm standard length, Lorestan, Dez or Karkheh River drainage (no other locality data); CMNFI 1979-0271, , mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0272, , mm standard length, Lorestan, river at Nokhor (33º40-47'N, 48º28-45'E); CMNFI 1979-0273, 7, 66.7-137.6 mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0274, 3, 28.9-141.8 mm standard length, Lorestan, Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0276, , mm standard length, Lorestan, Kashkan River drainage (ca. 33º19'N, ca. 47º53'30"E); CMNFI 1979-0277, 2, 116.2-133.4 mm standard length, Lorestan, Kashkan River drainage (33º30'N, 47º59'30"E); CMNFI 1979-0278, 3, 93.5-114.7 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 1, 126.0 mm standard length, Lorestan, Khorramabad River 16 km from Nurabad (33º37'N, 48º18'E); CMNFI 1979-0279, 5, 115.6-155.8 mm standard length, Lorestan, Khorramabad River 16 km from Nurabad (33º37'N, 48º18'E); CMNFI 1979-0280, 3, 104.7-107.7 mm standard length, Lorestan, Kashkan River drainage (33º43-47'N, 48º12-15'E); CMNFI 1979-0282, 6, 110.3-130.3 mm standard length, Lorestan, Seymareh River drainage at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0283, 2, 113.7-125.0 mm standard length, Kermanshahan, Qareh Su drainage (34º21'N, 47º07'E); CMNFI 1979-0285, 3, 125.5-148.0 mm standard length, Kermanshahan, Qareh Su drainage (34º26'N, 46º37'E); CMNFI 1979-0286, , mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0287, 2, 128.2-136.1 mm standard length, Kermanshahan, Chashmeh Javari 2 km from Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0288, 62, 37.6-153.7 mm standard length, Ilam and Poshtkuh, Gangir River at Sarab Ewan (33º50'N, 46º18'E); CMNFI 1979-0289, , mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, , mm standard length, Kermanshahan, Diyala River drainage at Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, , mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0306, , mm standard length, Kerman, qanat on road to Baft (29º13'N, 54º33'E); CMNFI 1979-0307, 5, 50.9-73.4 mm standard length, Kerman, river at Sartal 6 km from Baft (ca. 29º17'N, ca. 56º38'E); CMNFI 1979-0308, 67, 20.5-246.9 mm standard length, Kerman, river 44 km from Baft (29º02'N, 56º50'E); CMNFI 1979-0309, , mm standard length, Kerman, Fahraj River at Azizabad (28º57'N, 58º42'E); CMNFI 1979-0315, 2, 53.5-65.5 mm standard length, Baluchestan, Bampur River 2 km from Karevandar (27º51'N, 60º46'E); CMNFI 1979-0315, 34?, 53.7-85.1 mm standard length, (); note two collections? CMNFI 1979-0337, , mm standard length, Baluchestan, stream near Kanowak (ca. 28º40'N, ca. 60º48'E); CMNFI 1979-0341, 14, 27.2-75.9 mm standard length, Kerman, Tahrud west of Bam (29º23'N, 57º52'E); CMNFI 1979-0343, , mm standard length, Fars, lake near Deh Bid (ca. 30º32'N, ca. 52º49'E); CMNFI 1979-0411, 7, 42.2-76.5 mm standard length, Hormozgan, Minab River past Rudan (27º24'N, 57º12'E); CMNFI 1979-0419, 1, 62.2 mm standard length, Fars, stream 7 km from Rostaq (28º29'N, 55º01'E); CMNFI 1979-0420, 6, 57.1-150.6 mm standard length, Fars, Rudbar River at Bahregan (30º11'N, 52º03'E); CMNFI 1979-0422, , mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Yasuj valley (30º36'N, 51º34'E); CMNFI 1979-0424, , mm standard length, Fars, stream on Yasuj to Nurabad road (30º18'N, 51º30'30"E); CMNFI 1979-0425, , mm standard length, Fars, Haft Barm-e Kudian (29º49'N, 52º02'E); CMNFI 1979-0426, , mm standard length, Esfahan, qanat at Abbasabad-Natanz (33º36'N, 51º49'E); CMNFI 1979-0458, 2, 90.7-108.4 mm standard length, Markazi, Khar River 6 km north of Ab-Garm (35º47'N, 49º20'E); CMNFI 1979-0460, 3, 54.4-65.0 mm standard length, Hamadan, stream 16 km south of Asadabad (34º39'N, 48º05'E); CMNFI 1979-0462, , mm standard length, Markazi, Mazdaqan River (35º06'30"N, 49º40'30"E); CMNFI 1979-0466, , mm standard length, Esfahan, qanat at Meymeh (33º27'N, 51º10'E); CMNFI 1979-0484, , mm standard length, Khorasan, stream 22 km west from Bojnurd (37º28'N, 56º44'E); CMNFI 1979-0497, 3, 49.8-113.0 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0497, 7, 102.2-132.0 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0499, , mm standard length, Fars, ditch 32 km from Kor River bridge (30º04'30"N, 52º36'E); CMNFI 1979-0501, 6, 34.1-110.9 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0502, , mm standard length, Fars, Haft Barm-e Kudian (29º49'N, 52º02'E); CMNFI 1993-0126, , mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 2007-0030, , mm standard length, Baluchestan, stream near Eskelabad (28º35'N, 60º48'E); CMNFI 2007-0031, , mm standard length, Baluchestan, headwaters of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0037, , mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, , mm standard length, Kerman, Mehtiabad qanat (29º06'N, 58º21'E); CMNFI 2007-0039, , mm standard length, Kerman, Tahrud River (ca. 29º23'N, ca. 57º53'E); CMNFI 2007-0040, , mm standard length, Kerman, Qahariz qanat at Jupar (30º04'N, 57º08'E); CMNFI 2007-0041, , mm standard length, Kerman, qanat at Baghin (30º12'N, 56º48'E); CMNFI 2007-0042, , mm standard length, Kerman, qanat at Negar (29º52'N, 56º50'E); CMNFI 2007-0043, , mm standard length, Kerman, qanat at Emamzadeh Sultan (ca. 29º40'N, ca. 56º45'E); CMNFI 2007-0044, , mm standard length, Kerman, Qal'eh-ye Askar stream (ca. 29º28'N, ca. 56º38'E); CMNFI 2007-0045, , mm standard length, Kerman, Kharan River drainage at Baft (29º14'N, 56º38'E); CMNFI 2007-0047, , mm standard length, Kerman, qanat at Hoshun (29º14'N, 56º19'E); CMNFI 2007-0048, , mm standard length, Kerman, qanat at Hasanabad (ca. 28º50'N, ca. 55º50'E); CMNFI 2007-0049, , mm standard length, Hormozgan, upper Kol River basin at Hajjiabad (ca. 28º19'N, ca. 55º55'E); CMNFI 2007-0063, , mm standard length, Fars Mand River tributary outside Jahrom (28º36'N, 53º37'E); CMNFI 2007-0065, , mm standard length, Fars, Barm-e Dalak (ca. 29º35'N, ca. 52º38'E); CMNFI 2007-0066, , mm standard length, Fars, qanat under Sa'adi's Tomb, Shiraz (29º37'N, 52º35'E); CMNFI 2007-0067, , mm standard length, Fars, Sivan River (ca. 30º02'N, ca. 52º57'E); CMNFI 2007-0068, 5, 59.0-89.6 mm standard length, Fars, qanat 4 km south of Abarqu (ca. 31º07'N, ca. 53º14'E); CMNFI 2007-0069, , mm standard length, Yazd, qanat at Zarej (ca. 31º58'N, ca. 54º17'E); CMNFI 2007-0070, , mm standard length, Yazd, qanat at Ardakan, (32º19'N, 53º59'E); CMNFI 2007-0073, , mm standard length, Esfahan, Zayandeh River at Tanderan (32º47'N, 51º02'E); CMNFI 2007-0075, , mm standard length, Hamadan, Malayer River south of Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0076, , mm standard length, Markazi, Malekabad qanat (34º05'N, 49º53'E); CMNFI 2007-0083, , mm standard length, Azarbayjan-e Khavari, Qaranqu River basin west of Sar Eskand Khan (ca. 37º25'N, ca. 46º55'E); CMNFI 2007-0084, , mm standard length, Azarbayjan-e Khavari, Talkheh River basin west of Sarab (ca. 37º56'N, ca. 47º19'E); CMNFI 2007-0091, , mm standard length, Azarbayjan-e Khavari, Zilber Chay basin west of Marand (38º30'N, 45º23'E); CMNFI 2007-0100, , mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0108, , mm standard length, Kordestan, Qeshlaq River basin north of Sanandaj (ca. 35º33'N, ca. 47º08'E); CMNFI 2007-0109, , mm standard length, Kordestan, Qeshlaq River basin south of Sanandaj (ca. 35º16'N, ca. 47º01'E); CMNFI 2007-0110, , mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0115, , mm standard length, Kermanshahan, Qareh Su basin north of Kermanshah (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, , mm standard length, Kermanshahan, Gav Masiab River basin west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, , mm standard length, Kermnashahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0117, , mm standard length, Kermnashahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0122, , mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); USNM 200308, 2, 37.5-47.3 mm standard length, Lorestan, Ab-e Khorramabad (33º30'N, 48º13'E); ? damascina USNM 205933, 5, 97.5-142.4 mm standard length, Baluchestan, Karavandar Creek (no other locality data); ? damscina USNM acc. 303854, 14, 29.1-44.6 mm standard length, Fars, Lake Arzhan (29º36'N, 51º59'E). ? damscina USNM acc. 303854, 4, 55.7-99.0 mm standard length, Fars, pool east of Sangkar (29º48'N, 53º29'E); ?damascina

Capoeta aculeata
(Valenciennes, 1844)

  
                Left pharyngeal arch

Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi

Common names

shum (= unlucky or inauspicious, possible meaning), سياه ماهي (= siah mahi), زرده پر (= zardehpar), siah mahi aculeata.

Systematics

Chondrostoma aculeatum was originally described from "eaux douces de la Perse".

Scaphiodon macrolepis Heckel, 1847 described from the "Confluenten des Araxes bei Persepolis" (probably the Pulvar (= Sivan) River, Fars near Persepolis) and Varicorhinus bergi Derzhavin, 1929 described in Latin from "Keredsh flumen propea Teherane, Persia septentrionalis" (Karaj River near Tehran, northern Iran) are synonyms.

Six syntypes (MNHN 2357) of Chondrostoma aculeatum in poor condition are stored in the Muséum national d'Histoire naturelle, Paris (Bertin and Estève, 1948; Coad and Krupp, 1994). They measure 86-179 mm standard length (Coad and Krupp, 1994) or 105-210 mm total length (Bertin and Estève, 1948). The largest specimen is designated as the lectotype.

Two syntypes of Scaphiodon macrolepis are in the Naturhistorisches Museum Wien under NMW 55896 and measure 100-175 mm standard length (Kähsbauer, 1964). Two other fish are marked as syntypes under NMW 51653 and are from Persepolis collected by Th. Kotschy. The catalogue in Vienna lists 4 fish and the 1997 card index agrees these 4 fish are the syntypes.

Types of Varicorhinus bergi are unknown (Eschmeyer et al. (1996)).

Berg (1949) considers both aculeata and macrolepis as distinct species although very close, the latter distinguished from the former by a deeper body and a shorter head. Karaman (1969) and Bianco and Banarescu (1982) place both aculeata and macrolepis in Capoeta capoeta; Karaman does suggest that macrolepis could belong in aculeata. Saadati (1977) considers aculeatus not more than subspecifically distinct from macrolepis, not realising the former has priority.

Key characters

This species differs from all others in the genus Capoeta in the lower number of lateral line (93% of 314 fish had range of 39-48) and caudal peduncle scales (90% of 303 fish had a range of 16-20). Capoeta capoeta, a related species, usually has 54 or more lateral line scales and 20 or more caudal peduncle scales.

Morphology

Dorsal fin with 3-5, modally 4, unbranched and 7-9, modally 8, branched rays. The last dorsal fin unbranched ray is thickened and serrated, the denticles being long and narrowly spaced but not strongly developed. Distally this spiny ray is flexible. Smaller fish have proportionately larger and more extensive denticles than larger fish. The extent of denticles from the base distally varies between about two-thirds and three-quarters. Anal fin with 3 unbranched and 5-6, modally 5, branched rays, pectoral fin with 14-21 branched rays, and pelvic fin with 7-10 branched rays.

Lateral line scales 36-52. Caudal peduncle scales 13-23. Scale shape is squarish with shallowly rounded to straight dorsal and ventral margins, sharp corners anteriorly, and a large to moderate central protuberance on the anterior margin. Radii are most numerous on the posterior field but even there are few, relatively few laterally and few anteriorly. Circuli are very fine but break into coarser "bubbles" on the posterior field. The focus is subcentral anterior. The pelvic fin axillary scale varies greatly in size.

The mouth is slightly arched or even straight in ventral view. The horny edge to the lower jaw is usually well-developed but may be lost in preserved specimens. Gill rakers number 16-25 and are short, reaching past the first or second raker when appressed. Rakers are thick and usually hooked at their tips. Pharyngeal teeth are modally 2,3,4-4,3,2 (in 10 fish). Major row teeth are spatulate with a wide crown in large fish. Total vertebrae number 39-44. The gut is extremely elongate with numerous anterior and posterior coils.

Meristic characters in Iranian fish are: dorsal fin branched rays 7(50), 8(255) or 9(4); anal fin branched rays 5(177) or 6(1); pectoral fin branched rays 14(2), 15(3), 16(12), 17(52), 18(123), 19(79), 20(27) or 21(6); pelvic fin branched rays 7(23), 8(183), 9(102) or 10(6); lateral line scales 36(1), 37(5), 38(8), 39(15), 40(25), 41(48), 42(56), 43(38), 44(40), 45(23), 46(22), 47(15), 48(10), 49(1), 50(4), 51(2) or 52(1); scales around the caudal peduncle 13(1), 15(5), 16(48), 17(52), 18(73), 19(64), 20(37), 21(12), 22(5) or 23(6); total gill rakers 16(3), 17(13), 18(40), 19(49), 20(62), 21(43), 22(43), 23(27), 24(16) or 25(6); and total vertebrae 39(1), 40(26), 41(90), 42(103) or 44(16).

Sexual dimorphism

Males have moderately large tubercles on the anal fin rays following the ray branching (2-4 tubercles on last 4 branched anal rays), small tubercles on the lowest caudal fin ray, very fine tubercles on top of the head, larger tubercles on the side of the head, largest on the snout below the eye and nostril as far as the mouth, connecting across the snout, and numbering 1-5 moderately large tubercles on flank scales variously arranged on each scale and best developed on the posterior part of the body.

Colour

The back is almost entirely black to green-brown or olive-green, the upper flank is brownish, the belly and lower flank are yellow up to the lateral line, only the belly centre being white. The flanks are generally silvery in live fish. Some fish have small black spots on the sides and fins. Preserved fish have pigment on the posterior, exposed margin and so are outlined on the flank. The sides of the head are golden-brown. Flank spots may be in 5 longitudinal rows above, and 2 rows below, the lateral line. Some populations have fish with spots and mottles on the body and fins but these are probably occasioned by a parasitic infestation. Fins are often reddish-brown to pink although pelvic and anal fins may be yellowish-green and the dorsal and caudal fins very light to hyaline. Preserved fish have pigment on the rays and membranes of fins without any distinctive pattern. The dorsal and caudal fins are darker than the lower fins. The iris is golden to orange. The peritoneum is black.

Size

Reaches 23.4 cm standard length.

Distribution

This species is found in the Tigris River, Namak Lake, Dasht-e Kavir, Kerman-Na'in, Esfahan, Kor River basins (Rainboth, 1981; Bianco and Banarescu, 1982; Ghorbani Chafi, 2000). Abdoli (2000) maps this species from the Kerman-Na'in basin generally; the upper Kal Shur, Jajarm and Jovein rivers in the Dasht-e Kavir basin; the middle and upper Shur and Abhar, Qareh Chai and Qom rivers in the Namak Lake basin; the Zayandeh and Shur rivers in the Esfahan basin; the Jarrahi and Marun, upper Karun and Khersan, Dez, Karkheh, Simarreh and Kashkan rivers in the Tigris River basin. Raissy et al. (2010) record it from the Armand River in Chaharmahal va Bakhtiari Province. K. Abbasi records it from the Gamasiab River (see photograph above).

Zoogeography

Saadati (1977) suggests that this species moved eastward to basins on the plateau during more pluvial periods from the Tigris River basin. See also above under genus.

Habitat

Unknown in detail. Abbasi et al. (2009) in their study of wetlands in Hamadan Province found this species was the third most dominant out of 23 species at 11.7%.

Age and growth

Unknown.

Food

Gut contents include filamentous algae, plant fragments and diatoms with large amounts of sand. This species has been seen turning belly up to feed (field notes for specimens from Jajarm, Khorasan).

Reproduction

Reproduction has not been studied in this species. Specimens from the Khorramabad River contained eggs 1.5 mm in diameter on 6 July and some seemed to be reabsorbing eggs. Spawning presumably takes place in late spring and summer.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Dactylogyrus lenkorani, Gyrodactylus sp. and Myxobolus sp. Masoumian et al. (2007) record the myxosporean parasite Myxobolus cristatus from this species in the Zayandeh River. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus chramuli, D. lenkorani and D. gracilis in the Zayandeh River. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea and Trichodina sp. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species. Raissy et al. (2010) found ichthyophthiriasis (infection with Ichthyophthirius multifilis - ich or white spot disease), which cause epizootics in wild and cultured fishes, in fish from the Armand River in Chaharmahal va Bakhtiari Province.

Economic importance

None.

Conservation

This species is widely distributed in Iran and does not appear to be in need of conservation but its biology and habitat requirements are unknown. Kamali-Far et al.(2009) have used carp pituitary extract in an attempt to induce spawning in this species. Hatchery production could then be used to supplement natural stocks. However, the attempt was unsuccessful. Note that the identity of the species used in this study needs verification judging from the photograph in the paper.

Further work

The biology of this species needs study as does its habitat requirements and conservation needs.

Sources

Type material: See above, Chondrostoma aculeatum (MNHN 2357).

Iranian material: CMNFI 1979-0025, 2, 65.3-68.1 mm standard length, Fars, Kor River near Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0059, 155, 22.9-67.4 mm standard length, Fars, Pulvar River 8 km south of Sivand (30º01'30"N, 52º57'E); CMNFI 1979-0061, 6, 28.6-64.9 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0069, 1, 28.7 mm standard length, Fars, qanat at Naqsh-e Rostam (29º59'30"N, 52º54'E); CMNFI 1979-0070, 16, 25.9-60.0 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0090, 2, 153.6-160.5 mm standard length, Esfahan, Gav Khuni (ca. 32º21'N, ca. 52º49'E); CMNFI 1979-0116, 49, 24.3-52.1 mm standard length, Fars, Kor River near Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0117, 14, 34.4-44.1 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 1979-0252, 3, ?, mm standard length, Markazi, jube at Baqerabad (34º55'N, 50º50'E); CMNFI 1979-0253, 5, 40.4-103.7 mm standard length, Markazi, stream in Qareh Chay drainage (34º52'N, 50º49'E); CMNFI 1979-0270, 1, 121.8 mm standard length, Lorestan, Kashkan River draiangae outside Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0271, 1, 52.1 mm standard length, Lorestan, stream in Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 28, 51.4-104.5 mm standard length, Lorestan, stream in Kashkan River drainage near Khorramabad (33º26'N, 48º19'E); CMNFI 1979-0274, 6, 20.6-59.2 mm standard length, Lorestan, stream in Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0275, 1, 50.9 mm standard length, Lorestan, Kashkan River 2 km from Ma'mulan (33º25'N, 47º58'E); CMNFI 1979-0279, 18, 41.1-129.9 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0282, 7, 99.2-130.8 mm standard length, Lorestan, river at Nurabad (34º05'N, 47º58'E); CMNFI 1979-0283, 2, 125.2-186.3 mm standard length, Kermanshahan, Qareh Su near Kermanshah (34º21'N, 47º07'E); CMNFI 1979-0343, 1, 146.6 mm standard length, Fars, lake near Deh Bid (ca. 30º32'N, ca. 52º49'E); check ID? CMNFI 1979-0365, 1, 25.0 mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30"E); CMNFI 1979-0396, 9, 32.5-58.7 mm standard length, Khuzestan, Kheyrabad River 20 km from Behbehan (30º32'N, 50º23'30"E); CMNFI 1979-0427, 2, 100.5-112.2 mm standard length, Markazi, Cheshmeh Fin at Fin (33º57'N, 51º24'E); checkID? CMNFI 1979-0428, 17, 25.9-104.5 mm standard length, Markazi, stream 3 km south of Sen Sen (34º13'N, 51º16'E); checkID? CMNFI 1979-0458, 9, 48.5-117.8 mm standard length, Markazi, Khar River 6 km north of Ab-garm (35º47'N, 49º20'E); CMNFI 1979-0460, 1, 77.6 mm standard length, Hamadan, stream 16 km south of Asadabad (34º38'N, 48º03'E); checkID? CMNFI 1979-0463, 8, 97.9-135.3 mm standard length, Markazi, Qareh Chay (34º53'N, 50º24'E); checkID? CMNFI 1979-0464, 1, 74.2 mm standard length, Markazi, qanat at Kheyrabad (34º08'N, 50º00'E); CMNFI 1979-0465, 18, 35.7-58.3 mm standard length, Markazi, Qom River (34º18'30"N, 50º32'E); CMNFI 1979-0500, 2, 92.4-98.6 mm standard length, Fars, Pulvar River near Naqsh-e Rostam (29º59'N, 52º54'E); checkID? CMNFI 1980-0156, 27, ? mm standard length, Markazi, Karaj River (35º47'N, 50º58'E); CMNFI 1993-0154, 1, mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); checkID? CMNFI 1993-0156, 1, mm standard length, Markazi, Sharra River (34º03'N, 49º21'E); checkID? CMNFI 2007-0006, 9, 59.9-127.2 mm standard length, Khorasan, spring in Qareh Su basin south of Garmeh (ca. 36º58'N, ca. 56º15'E); CMNFI 2007-0007, 8, 59.4-79.3 mm standard length, Khorasan, stream supplemented by qanats, Kal-e Tangeh (ca. 36º59'N, ca. 56º29'E); CMNFI 2007-0008, 2, 72.1-84.3 mm standard length, Khorasan, qanat at Jajarm (36º57'N, 56º23'E); CMNFI 2007-0009, 18, 35.9-108.1 mm standard length, Khorasan, qanat at Amirabad (ca. 36º31'N, ca. 56º45'E); CMNFI 2007-0010, 11, 80.8-123.1 mm standard length, Khorasan, qanat at Haresabad (36º07'N, 57º37'E); CMNFI 2007-0011, 12, 34.1-85.4 mm standard length, Khorasan, Kalshur River south of Neyshabur (36º05'N, 58º43'E); CMNFI 2007-0071, 10, 70.4-156.9 mm standard length, Esfahan, qanat at Mohammadiyeh, Na'in (32º51'N, 53º06'E); CMNFI 2007-0074, 29, 50.6-100.7 mm standard length, Markazi, Qareh Chai west of Arak (34º03'N, 49º21'E); CMNFI 2007-0075, 16, 29.3-152.5 mm standard length, Hamadan, Hamadan, Malayer River 5 km from Malayer (ca. 34º17'N, ca. 48º47'E); CMNFI 2007-0076, 5, 56.1-97.4 mm standard length, Markazi, Malekabad qanat east of Arak (34º05'N, 49º53'E); CMNFI 2007-0078, 8, 37.6-102.8 mm standard length, Markazi, Qom River (ca. 34º18'N, ca. 50º32'E); CMNFI 2007-0117, ?, mm standard length, Kermanshahan, Gav Masiab River basin near Sahneh (ca. 34º24'N, ca. 47º40'E); CMNFI 2007-0119, ?, mm standard length, Kermanshahan, Gav Masiab River basin near Kangavar (ca. 34º31'N, ca. 48º03'E); CMNFI 2007-0120, 15, 29.0-165.5 mm standard length, Hamadan, Ab Chay near Hamadan (ca. 34º49'N, ca. 48º29'E); CMNFI 2007-0122, 12, 35.0-77.6 mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); BM(NH) 1934.10.29:2, 1, 84.0 mm standard length, Markazi, Tehran (no other locality data); BM(NH) 1958.11.7:1-6, 6, 25.6-89.9, Khorasan, Jajarm (36º57'N, 56º23'E); BM(NH) 1975.1.17:255-258, 4, 103.0-161.0 mm standard length, Esfahan, Esfahan (no other locality data); MNHN 1960-611, 2, 127.0-144.0 mm standard length, Markazi, Jajrud east of Tehran (ca. 35º45'N, ca. 51º42'E) USNM 205932, 3, 78.5-159.4 mm standard length, Markazi, stream southwest of Tehran (35º34'N, 51º03'E); ZMH 5905, 2, 57.0-70.0 mm standard length, ?, Jafar Abad qanat (?); ZSM 25703, 1, 76.3 mm standard length, ?, Khorramabad River (no other locality data).

Capoeta barroisi
Lortet, 1894

Common names

siah mahi-ye Dasht-e Arzhani (Arzhan Plain black fish).

[tela barroisi in Arabic; spotted barb, Tigris barb].

Systematics

Subspecies are Capoeta barroisi persica Karaman, 1969 described from "See Zariwar, Mariwan, 120 km westlich v. Sannadaj" (Lake Zaribar near Marivan, Kordestan in the Tigris River basin) and Capoeta barroisi mandica Bianco and Banarescu, 1982 from the "Mand River near Dasht-e-Arzhan" of Fars Province. Krupp (1985c) considers both these to be synonyms of the nominal subspecies, C. b. barroisi.

The subspecies persica is distinguished from the type subspecies by having a more horseshoe-shaped mouth, 8 branched dorsal fin rays, 18 gill rakers, blackish pectoral, pelvic and anal fins, few but very large black spots on the body, a shorter anal fin and a longer pectoral fin, and a deep body, based on a single specimen. Krupp (1985c) considers the characters of mouth form and colour to fall within the range of the nominal subspecies (and by implication the other characters too). Özuluğ and Freyhof (2008) found it difficult to reach a conclusion on the taxonomic status of this subspecies on the basis of a single specimen which could be abnormal.

C. barroisi mandica differs from the type subspecies (C. barroisi barroisi) and C. b. persica in number of scales (61-68 in mandica (58-68 in types examined by me), 69-82 in barroisi, 78-79 in persica), number of gill rakers (21-24 in mandica (22-27 in types examined by me and apparently number is related to size of fish), 27-31 in barroisi, 18 in persica), from barroisi in having usually 8 branched dorsal fin rays (barroisi has 9 but persica also has 8), and from persica by a straight mouth (also straight or transverse in barroisi, arched in persica). Krupp (1985c) considers the scale counts to be within the lower range of the nominal subspecies, gill raker counts and mouth position do not differ from the nominal subspecies, and the dorsal fin ray count of 8 is seen in the subspecies mandica. Krupp observes that meristic and morphometric characters are extremely variable in widely distributed Capoeta species. Özuluğ and Freyhof (2008) examined 5 juvenile specimens from the Mand River and consider the subspecies mandica to be a valid species. Widespread taxa like Capoeta species are prime candidates for  molecular analyses which might help resolve conflicting views on - a single widespread, variable species versus several distinct species.

Berg (1949) considers this species to be close to C. damascina, differing by having a stronger spine in the dorsal fin, hardly an invariant character. Saadati (1977) considers that C. barroisi of Karaman is in fact C. damascina. Turan (2008) using mt 16S rDNA concluded on this evidence that C. barroisi was a subspecies of C. damascina although other genetic markers should be used for a more reliable assessment.

Syntypes of Capoeta barroisi are in the Musée Guimet d'Histoire Naturelle, Lyon (MGHN 3492, 316 mm standard length, from the Orontes near Antakya in Turkey collected by E. Chantre and MGHN 3493, 278 mm standard length, from Buhairat Hims in Syria collected by Th. Barrois) (Krupp, 1985c).

The holotype of C. barroisi persica is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH H4119, 185.2 mm standard length, Daryacheh-ye Zaribar, 35°32'N, 46°08'E, IV. 1968, W. Nümann (Bianco and Banarescu, 1982; Krupp, 1985c; examined and measured by me).

The holotype of C. b. mandica, 106.9 mm standard length, is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (IZA 7890), with 95 paratypes from the same locality in IZA 7891 (now numbering 84 fish measuring 34.2-84.9 mm standard length) and 5 paratypes in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB 3123), these 100 specimens having a standard length of 34-86 mm. Six paratypes of mandica are in the Canadian Museum of Nature, Ottawa under CMNFI 1982-0366 (from IZA 7891).?lengths

Key characters

The dorsal fin branched ray count of usually 9 rays is characteristic for the type subspecies but not nominal Iranian ones. Gill raker counts, a head length greater than the dorsal fin spine length and the irregular brownish markings on a silvery-white body are also distinctive.

Morphology

Dorsal fin with 3-4 unbranched and 8-10 branched rays, usually 9 in the type subspecies but 8 in the subspecies mandica, anal fin with 3 unbranched and 5 branched rays. Pectoral fin branched rays 15-16, pelvic fin rays 7-8. Lateral line scales 61-82, 58-68 in the subspecies mandica. Scales on the belly in front of the pelvic fins are small and may not be imbricate. Pharyngeal teeth 2,3,4-4,3,2. Gill rakers 18-31, reaching the second adjacent raker when appressed. The last dorsal fin unbranched ray is very strong, but narrows distally, and bears large denticles or serrations on three-quarters of its length. The snout in the holotype of the mandica subspecies has a depression in front of the nostrils.

ZMH6086 D8, A5 P15, V7, ll 58, gr 24, flanks speckled dorsally, mouth a gentle arch almost straight, large denticles in damaged D spine.

Sexual dimorphism

Tubercles in males are found from eye to eye around the snout with fine tubercles sparse on the top of the head. Most flank scales have a single, centrally-placed tubercle as do scales on the caudal peduncle. There is a single row of tubercles on the last three anal fin rays. There are some weak tubercles on the side of the head.

Colour

There are numerous, small, distinctive brown to black spots on the head, flank and dorsal and anal fins. The back and upper flank are silvery-white with slate to violet overtones and the belly is yellowish. The holotype of the subspecies mandica has light specking and mottling on the upper flank and back. All fins have some speckling on the rays and membranes but no clear rows of spots. Smaller fish (paratypes) have darker and bigger speckles than the holotype which extend lower on the flank.

Size

Reaches 31.6 cm standard length.

Distribution

The subspecies C. barroisi barroisi is found in the Ceyhan, Orontes, Quwayq and Tigris-Euphrates basins, C. b. persica only in Lake Marivan or Zaribar of the Tigris River basin of Iran, and C. b. mandica in the Mand River of the Gulf basin. Krupp (1985c) includes Iranian Gulf drainages for the type subspecies. Abdoli (2000) has Jarrahi and lower Karun in the Tigris River basin and the middle and lower Hilleh, lower Mand, and lower Dasht-e Palang rivers in the Gulf basin.

Zoogeography

Taxa in Iran are remote from the type locality of this species and may be indicative of the links between the Levantine fauna and the former tributaries of the Tigris-Euphrates basin in southern Iran. See also above under genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Nothing is known of biology and a conservation assessment cannot be made. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs study as a prerequisite for a conservation assessment. The relationships of nominal Iranian taxa to material from the type locality need further study using molecular techniques.

Sources

Type material: Capoeta barroisi mandica (IZA 7890, 7891, CMNFI 1982-0366 (from IZA 7891)) and C. barroisi persica (ZMH H4119).

Iranian material: ZMH 6086, 1, 73.6 mm standard length, Fars, Shur Fluβ, zufluβ von Mand-Fluβ (= C. b. mandica) P. Bianco ?check this against his paper.

Comparative material: BMNH 1974.2.22:1853-1856, 3, 48.9-60.2 mm standard length, Iraq, Kaliasan near Sulaymaniyah

Capoeta buhsei
Kessler, 1877

Gharechay, Namak Lake basin, May 2008, courtesy of K. Abbasi
Gharechay, Namak Lake basin, May 2008, courtesy of K. Abbasi

Common names

shamshiri (= sword-like), mahi sibili (= moustached fish, from Karaj Lake).

Systematics

Varicorhinus nikolskii Derzhavin, 1929 described in Latin from the "Keredsh flumen" (= Karaj River near Tehran) is a synonym. Saadati (1977) places Capoeta buhsei in Capoeta damascina.

The 2 syntypes of Capoeta Buhsei, 200.7-211.4 mm standard length, are in the Zoological Institute, St. Petersburg (ZISP 2330) and were collected "iz Persii" (= from Persia) by Dr. Buhse in 1849. The 11 syntypes of Varicorhinus nikolskii have not been located (Eschmeyer et al., 1996).

Key characters

This species is distinguished by its low total gill raker count of 9-17, mean 12.4, modes 12 and 13, lower arch rakers 7-10 (cf. C. damascina which has 17-25 total rakers, mean based on ? specimens), the absence of a keel in front of the dorsal fin, the mouth structure, and by a very weak, unserrated or barely serrated dorsal fin spine in large fish (cf. C. damascina).

Morphology

Dorsal fin unbranched rays 3-4, branched rays 7-9, anal fin unbranched rays 3, branched rays 5., pectoral fin branched rays 14-19, and pelvic fin branched rays 7-9. Lateral line scales 72-99. Scales are found on the back and on the belly. A pelvic axillary scale is present. Scales have parallel dorsal and ventral margins, a rounded posterior margin and an anterior margin with a rounded central protuberance. Radii are found on all fields including a few long and curved ones on the lateral fields. The focus is subcentral anterior and circuli are numerous and fine. Gill rakers 9-19 in literature (but see below), including some counts probably for the lower arm only, and reach the second raker below when appressed but only the next raker in small fish. Pharyngeal teeth in the main row are spatulate, the crowns flat, narrow and curved. Tooth counts are 2,3,4 or 5-5 or 4,3,2. The fifth tooth in either row is small and variably present. This may be size related although the fish examined here were all relatively small and showed no clear trend. The gut is elongate with several long coils.

The mouth is large and a shallow horseshoe-shape with the horny lower jaw layer weakly developed but the lower lip corners in particular fleshy and well-developed. The last unbranched dorsal fin ray is weak with the distal half thin and flexible. Denticles are found on the basal third to two-thirds or more of the ray, their extent and size variable but usually small, weak and less extensive in large fish, while larger and more extensive in the smallest fish. For fish 48.9-174.0 mm standard length extent of dorsal fin spine serrations in spine length is 0.3-0.8, mean 0.6. The lower lip is apparent and finely ridged. The upper lip and snout are covered with unculi which occur also over the head but more widely spaced out. The upper lip unculi are densely concentrated and are broader than other head ones. Unculi are also on the lower head surface and belly scales back to the pelvic fins, and on the anal, pectoral and pelvic fin rays and membranes.

Meristics are as follows: dorsal fin branched rays 8(35) or 9(3); anal fin branched rays 5(38); pectoral fin branched rays 14(1), 17(20), 18(11) or 19(6); pelvic fin branched rays 7(1), 8(5), 9(31); lateral line scales 72(2), 73(1), 75(3), 76(4), 77(2), 78(1), 79(7), 80(4), 81(5), 82(2), 83(2), 84(2), 86(2) or 91(1); total gill rakers 9(2), 10(2), 11(4), 12(12), 13(11), 14(4), 15(2) or 17(1); pharyngeal teeth 2,3,4,-4,3,2(7), 2,3,4-5,3,2(6), 2,3,5-4,3,2(5), 2,3,5-5,3,2(1), 1,3,4-4,3,2 (1) or 2,3,4-4,3,1(1); and total vertebrae 43(1) and 44(6) (USNM 20593 and the syntypes).

Sexual dimorphism

One male specimen measuring 94.6 mm standard length bears large tubercles on anal fin rays, fine tubercles scattered on the head, on the back and upper flanks one tubercle per scale at the scale centre but not on every scale, all along the lateral line at one tubercle per scale, and below the lateral line only in the area above the anal fin.

Colour

Overall colour is brownish in preservative without spots or any distinctive markings. The back is dark. The peritoneum is dark brown to black in preserved fish.

Size

Reaches 25.7 cm.

Distribution

This species is endemic to the Namak Lake basin of Iran (Derzhavin, 1929; Wossughi, 1978; Abdoli, 2000). Abdoli (2000) questionably maps it from the Esfahan basin. A report from Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994) is presumably a mis-identification and records from springs of Kul River basin near Darab in the Hormuz basin (Bianco and Banarescu (1982) and the Hamun-e Jaz Murian basin (Vossoughi, 1998) are also questionable.

Zoogeography

An endemic of an interior Iranian basin, its zoogeographical relationships to other Capoeta have not been resolved. See also above under genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Gut contents include aquatic insect larvae and masses of filamentous algae, suggesting that aufwuchs is an important diet item.

Reproduction

Generally unknown but fish caught on 5 June measuring 121.3-132.6 mm standard length have small eggs, perhaps because this size of fish is not mature. A 174.0 mm standard length caught in January has larger eggs than those from the June fish. A male fish caught on 5 May and measuring 146.6 mm standard length has mature testes.

Parasites and predators

Williams et al. (1980) report the helminths Khawia armeniaca (a cestode) and Acanthocephalorhynchoides cholodkowskyi (an acanthocephalan) from this species in the Zayandeh River at Esfahan.

Economic importance

Unknown.

Conservation

The conservation status of this species has not been determined by field studies and assessments can only be done from museum collections.

Further work

The distribution and numbers of this species in the Namak Lake basin should be examined by field studies to determine the population status. This basin is mostly in Markazi (= Central) Province which contains Tehran and a very large human population with great demands on limited water resources. It is probably not under any immediate threat but is an Iranian endemic.

Sources

Type material: See above, Capoeta buhsei (ZISP 2330).

Iranian material: CMNFI 1970-0588, 19, 42.4-128.9 mm standard length, Markazi, Karaj Lake (35º57'N, 51º06'E); CMNFI 1979-0094, 2, 143.1-174.0 mm standard length, Markazi, Karaj Lake (35º57'N, 51º06'E); CMNFI 1979-0266, 2, 52.4-54.3 mm standard length, Esfahan, spring at Nowqan (ca. 33º10'N, ca. 50º05'E); CMNFI 1979-0458, 1, 94.2 mm standard length, Markazi, Khar River (35º47'N, 49º20'E); CMNFI 1979-0459, 2 ?check fish, only 1 in catalogue, 27.0-31.6 mm standard length, Hamadan, stream 2 km south of Razan (35º22'N, 49º02'E); CMNFI 1979-0461, 1, 54.1 mm standard length, Hamadan, qanat at Taveh (35º07'N, 49º02'E); CMNFI 1979-0495, 1, 42.5 mm standard length, Markzai, Nam River west of Firuzkuh (35º43'N, 52º40'E); CMNFI 1980-0154, 71, 12.0-34.9 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1980-0156, 27, 32.4-54.3 mm standard length, Markazi, Karaj River below village (35º47'N, 50º58'E); CMNFI 1993-0151, 1, 146.4 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º19'E); CMNFI 1993-0152, 2, 121.3-132.6 mm standard length, Markazi, Sharra River near Khosbijan (34º07'N, 49º23'E); CMNFI 1993-0153, 2, 104.3-138.9 mm standard length, Markazi, Sharra River near Emarat (33º52'N, 49º36'E); CMNFI 1993-0154, 1, 124.0 mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); CMNFI 2007-0074, 3, ? mm standard length, Markazi, Qareh Chay (34º03'N, 49º21'E); CMNFI 2007-0078, 5, ? mm standard length, Markazi, Qom River (ca. 34º18'N, ca. 50º32'E): check ID? CMNFI 2007-0079, 14, ? mm standard length, Zanjan, Abhar River basin (ca. 36º16'N, ca. 49º08'E); CMNFI 2007-0120, , mm standard length, Hamadan, Ab Chay (ca. 34º49'N, ca. 48º29'E); CMNFI 2007-0121, 3, 82.5-141.5 mm standard length, Hamadan, Qareh Su basin north of Razan (ca. 35º25'N, ca. 49º02'E); CMNFI 2007-0122, , mm standard length, Markazi, Khar River basin south of Takestan (ca. 35º56'N, ca. 49º30'E); USNM 20593, ?, ? mm standard length, (); ZMH 2632, 1, 148.2 mm standard length, Dojodje (); ZMH 2633, ?, ? mm standard length, above Latian ().

Capoeta capoeta
(Güldenstaedt, 1773)

Ghalechai, Lake Orumiyeh basin, October 2011, courtesy of K. Abbasi
Ghalechai, Lake Orumiyeh basin, October 2011, courtesy of K. Abbasi

Capoeta capoeta heratensis from Wikimedia Commons
Capoeta capoeta heratensis from Wikimedia Commons

Common names

tilkhos, سياه ماهي (= siah mahi, meaning black fish), sang lisak (= rock snail?); soru (= slippery) in the Dalaki and Shapur river basins.

[gara balig or Lankaran xramulyasi for C. c. gracilis, Kur xramulyasi for C. capoeta, both in Azerbaijan; khramulya, capoeta, kapuit, kaput (all apparently derived from local names in Georgia and Armenia, namely khramuli and kapweti); Lenkoranskaya khramulya or Lenkoran khramulya, Kurinskaya khramulya or Kura khramulya, Zakaspiiskaya khramulya or Transcaspian khramulya (also marinka is used locally for the Transcaspian khramulya subspecies but this is an error), Araksinskaya khramulya or Araks khramulya, all in Russian; Transcaucasian barb; khramulia; kersin handscherli at Aleppo, in Arabic].

Systematics

Cyprinus capoeta was originally described from Tbilisi, Georgia.

Cyprinus fundulus Güldenstaedt, 1787 from the Caspian Sea, Cyrus River (and Capoeta fundulus Valenciennes, 1842), Scaphiodon asmussii Keyserling, 1861 from "Warme Quelle bei Sultan Karaul, 8 Meilen nordöstlich von Herat" (now in Afghanistan, formerly in Persia), Scaphiodon gracilis Keyserling, 1861 from "Wasserleitung bei Gaes, einige Meilen von Isphahan", Scaphiodon heratensis Keyserling, 1861 described from the "Heri-Rud, ein Fluss bei Herat" (now in Afghanistan, formerly in Persia), Capoeta Hohenackeri Kessler, 1877 from Caucasia (probably lower Kura and Araks rivers, Azerbaijan), Capoeta (Scaphiodon) Steindachneri Kessler, 1872 and Capoeta Steindachneri var. platylepida Kessler, 1872 both from the Zeravshan River, Uzbekistan, and probably Capoeta gibbosa Nikol'skii, 1897 described in Latin as from "Bochsani in Persia orientale" are synonyms. Capöeta Guldenstädtii De Filippi in Tortonese, 1940 from "F. Arasse, Erzerum (Anatolia)" is Capoeta capoeta but it is a manuscript name and is not available (Tortonese, 1940; Eschmeyer et al., 1966); 2 syntypes are in the Istituto e Museo di Zoologia della R. Università di Torino (MZUT N.729).

Eschmeyer et al. (1996) have the date of Cyprinus capoeta as 1772; the type locality is Tiflis, Caspian Sea: they also have Cyprinus fundulus authored by Pallas, 1814 although Berg (1948-1949) has Güldenstädt as the author.

A hybrid of Capoeta capoeta heratensis and Schizothorax pelzami is reported from the northern Kopetdag in Turkmenistan (Starostin, 1936).

Capoeta hohenackeri Kessler, 1877 described from tributaries of the Kura and Aras rivers has a high lateral line scale count in the original description (78) and might be a mislabelled Capoeta tinca (Heckel, 1843) from Black Sea drainages of Georgia and Turkey rather than the Caspian Sea basin.

Capoeta capoeta gracilis is the subspecies of much of Iran and Capoeta capoeta heratensis (figure above) is the subspecies from the Tedzhen River basin (Berg, 1949). The former usually has one pair of barbels, the latter two pairs (but see below). Bianco and Banarescu (1982) limit C. c. gracilis to basins between the Safid River and the Atrak while C. c. capoeta is found in the Kura-Aras basin. Holčík and Jedlička (1994) consider that the two subspecies gracilis and heratensis do not exist but that the taxon C. capoeta exhibits clinal variation.

Bănărescu in Bănărescu (1999) limits C. capoeta gracilis to the Lake Orumiyeh basin and the Safid River in Iran (and the lower Kura River of Azerbaijan) while his C. capoeta aff. gracilis (an unnamed subspecies related to C. capoeta gracilis) is found along the rest of the Iranian Caspian shore. However his material was limited (and did not include any from Esfahan, the type locality of gracilis) and the analysis is based on lateral line scale counts only. Bănărescu in Bănărescu (1999) also states that C. capoeta sevangi de Filippi, 1865 is the subspecies of the Araxes River basin, presumably including Iran, distinguished from the type subspecies, C. capoeta capoeta of the Kura River basin, by having the dorsal fin margin straight or slightly convex as opposed to slightly to moderately notched. This character does not seem to be significant for such wide ranging and variable populations, which he admits in one case at least (Kura River at Mingechaur), show differences between samples from the same locality at different times.

Abdurakhmanov (1962) compares fish from the Kura River basin (presumably C. c. capoeta) with fish from the Lenkoranchai and Bilyashchai in Azerbaijan (C. c. gracilis) and finds that the latter have fewer dorsal fin rays on average, greater head length and depth, smaller eye, longer snout and postorbital distance, greater body depth and caudal peduncle depth, a shorter postdorsal distance, a shorter dorsal fin base, lesser dorsal fin height, a longer anal fin base, a greater pectoral-pelvic distance and a shorter pelvic-anal fin distance.

Dadikyan (1986) refers to Varicorhinus capoeta araxensis subsp. nov. from the Aras River basin in Armenia.

Günther (1899) points out that the considerable morphological variation shown by these fishes has resulted in numerous specific names and that it is difficult to assess them without a large comparative series and better information on localities. Berg (1948-1949) also indicates that the various subspecies are very close to each other and that their distributions are not clearly isolated.

C. c. heratensis shows major variations in body form, sometimes called morpha elata with a deep body and morpha elongata with a shallow and elongate body. These are not taxonomically significant but simply ecomorphs and all intermediates between the two extremes can be found. The deep-bodied form probably formed part of the fishes described as asmussii (Berg, 1948-1949). Reshetnikov and Shakirova (1993) list Capoeta heratensis as a full species.

Samaee et al. (2006) showed differences in morphometry between fish from six rivers along the Iranian Caspian shore with an overall assignment of individuals to a group of 88.6%. The morphometric data were mirrored by molecular data. Differences in morphometry were attributed to environmental and habitat conditions (temperature, turbidity, food availability and water depth and flow) but molecular data indicated a genetic basis, presumably through lack of gene flow between the river populations. Samaee et al. (2009) examined morphological variation with this species in the Shirud of the south Caspian Sea basin. There were no significant differences in meristic characters but morphometric characters varied and could be used to distinguish 5 groups. AnvariFar et al. (2010, 2011) compared fish from above and below the Shahid Rajaei Dam (built in 1995) and found the two populations to be morphologically different.

Records of Capoeta capoeta from the Tigris River basin at least are probably Capoeta damascina with low scale counts (F. Krupp, in litt., 1986).

Wossughi (1978) described, in a dissertation, a subspecies from the Namak Lake basin (from "Tschmeh Jafar Abad bei Araq") but this work may not be published in the sense of the International Code of Zoological Nomenclature (Ride et al., 1985). In any case, the holotype is Capoeta aculeata and the other material comprises 21 Leuciscus (= Squalius) cephalus orientalis and 4 Capoeta aculeata (F. Krupp, in litt., 1984). The type material, all female, is stored in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (holotype, 132 mm standard length under ZMH 5946, and 2 paratypes, 115-121 mm standard length, under ZMH 5947).

Bianco and Banarescu (1982) described C. capoeta intermedia from the Mand River in Fars but this is referred to C. damascina here (q.v.).

The types of Capoeta gibbosa are in the Zoological Institute, St. Petersburg (ZISP 11104) but have dried at some point. Their locality is given by Nikol'skii (1897) in Latin as "Bochsani in Persia orientali. 4.VII.96 (2)". This may be Bozjani at 35°48'N, 59°36'E. Berg (1949) considers that this nominal species is close to C. capoeta gracilis but is distinguished by body proportions (longer caudal peduncle and a longer head) but it is founded on only 2 specimens, hardly an adequate sample.

Types of Scaphiodon asmussii, S. gracilis and S. heratensis were not kept ? phrasing

Key characters

Berg (1948-1949) and Abdurakhmanov (1962) separate C. c. capoeta from C. capoeta gracilis, both of which may occur in Iran, by the following key:-

1(2). Dorsal fin emarginate above; lateral line scales usually 55-59; dorsal fin spine strong with numerous denticles; back behind occiput and particularly in front of the dorsal fin strongly compressed.....C. c. capoeta

2(1). Dorsal fin truncated in adults; lateral line scales usually 47-58; dorsal fin weak; back behind occiput not or only weakly compressed; radii on scales with minute recesses .....C. c. gracilis

Morphology

Dorsal fin unbranched rays 3-5, branched rays 7-10, usually 8-9; anal fin unbranched rays 2-4, branched rays 5; pectoral fin branched rays 15-20; and pelvic branched rays 7-9. Lateral line scales 46-70. In the subspecies gracilis the scales are said to be somewhat larger than in the type form, 47-58, mostly 48-50, and in heratensis the range is given as 50-60 by Berg (1948-1949). Scales are regularly arranged over the body. There is a pelvic axillary scale. Scales have a wavy anterior edge, few anterior and posterior radii and an almost central focus. Gill rakers 16-30, lower counts may refer to lower arm rakers only and total counts in the range 25-30 are probably more typical. Vertebrae 42-47. Pharyngeal teeth 2,3,4-4,3,2 or 2,3,5-4,3,2 with a hooked tip, spatulate below on posterior teeth while anterior teeth are conical. The last unbranched dorsal fin ray is strong with denticles along one half to two-thirds of its length (less strong in gracilis than in the type form). The number of barbels is variable - in fish from Uzbekistan 2 barbels (126 fish or 58.6%), 3 barbels (15 or 7.0%) or 4 barbels (74 or 34.4%)(Amanov, 1970). The subspecies heratensis is characterised by having 4 barbels but this is probably variable in Iran as in Uzbekistan. Six specimens from the Hari River basin of Iran all had 4 barbels.

Levin et al. (2005) found gracilis and heratensis (and steindachneri) to be oligovertebrate with 41-45 vertebrae, modes 42 to 44, compared to the multivertebrate type subspecies capoeta and sevangi with 45-48 vertebrae, mode 46. Morphometry and longevity also differ between these two groups and it was assumed they belong to different phyletic lines.

The Lake Sevan, Armenia subspecies (sevangi) has 2n=150 and is closer to "Barbus" than to African Varicorhinus, a genus in which Southwest Asian Capoeta were once placed (Krysanov, 1999). C. capoeta from the Safid River, the Shahrud in Rudbar and the Madarso River in Golestan National Park also have 2n=150, NF=230-234 (Pourali et al., 2000; Pourali Darestani et al., 2006). C. c. umbla from the Tigris River of Turkey had 2n=150, possibly hexaploid, with 43 meta-submetacentric chromosomes, 32 pairs of subtelo-acrocentric chromosomes with NF=236 (Kiliç Demirok and Ünlü, 2001).

Günther (1899) points out that this species shows considerable morphological variation, even in fish caught at the same place and time. The mouth can vary from straight to a gentle crescent to a distinct crescent, e.g. in three fish from the Nazlu Chai. There are also variations in dorsal fin spine development and the crown of the head can be flattened or convex.

Meristics for Iranian material: ?

Sexual dimorphism

The snout in males has 2-4 rows of tubercles and tubercles are present on scales and the rays of the anal fin. Abdurakhamanov (1962) reports that caudal peduncle length and lower caudal lobe length are longer in males while anal fin height, pelvic-anal fin distance, postorbital distance and interorbital width are greater in females.

Colour

The back is dark grey or green to brownish and the flanks light, silvery or silvery-grey, or yellowish. There may be several large black spots or blotches on the flank. The belly and lower head surface are pearly-white to dirty yellow. Scales are darkly pigmented. The operculum has a broad, yellow-gold spot. The iris is silvery, somewhat darker or yellow-golden above, or golden overall with traces of grey. The front of the dorsal fin and the margin of the caudal fin are black, and the rest of these fins are grey or yellowish-grey with some pink. The black margin to the caudal fin may be best developed on the upper and lower lobes compared to the posterior margin. The pectoral, pelvic and anal fins are grey with some pink or may be an overall pale pinkish. The peritoneum is black.

Size

Reaches 38 cm standard length and 3.5 kg (Amanov, 1970). This species reaches 43 cm in the Aras River basin of Iran (possibly C. capoeta sevangi, see above; A. Abdoli, pers. comm., 1995), 55 cm in Lake Sevan as C. capoeta sevangi (Bănărescu in Bănărescu, 1999) and 43.5 cm fork length and 1.23 kg in Çıır Lake, Turkey as C. capoeta capoeta.

Distribution

The type species is found in the Kura River basin of Azerbaijan with some Aras River basin fishes very similar. It is not known if the fishes from the Aras River basin in Iran belong to the type species or to gracilis. The subspecies Capoeta capoeta gracilis (known to Russian ichthyologists as the Lenkoran khramulya) was described from near Esfahan and is recognised as the one found over much of Iran including the Caspian Sea basin from the Astara to the Atrak including the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras, Tonekabon, Pol-e Rud and Safid rivers, teh Qezel Owzan, the Anzali Talab, the Dough Pishkamer, Tilabad, Zarrin-gol and Chelchal streams in the Gorgan River basin, and Gorgan Bay (Derzhavin, 1934; Bianco and Banarescu, 1982; Holčík and Oláh, 1992; Kiabi et al., 1994; Roshan Tabari, 1997; Shamsi et al., 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Abdoli and Naderi, 2009; Kazemian et al., 2009; Banagar et al., 2011 Shamekhi et al., 2012a, 2012b), the Lake Orumiyeh basin including the Urmi River or Shaher Chai, Ghalechai, Nazlu Chai and Talkheh, Tatavi and Zarrineh rivers (Günther, 1899; Abdoli, 2000; K. Abbasi, see photograph above) and Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), Marmisho Lake, west of Orumiyeh (Shiri et al., 2009; Ramin et al., 2012), the Dasht-e Kavir basin (Saadati, 1977), the Gulf basin in the Zohreh, Shapur, Dalaki, Helleh, Shur rivers and the upper and lower Mand River as Capoeta capoeta intermedia ((Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha (pers. comm., 1995; Abdoli, 2000), the Esfahan basin including the Dopolan River, Gav Khuni marsh (Keyserling, 1861), the Tigris River basin in the Regab River near Kermanshah, Nashad River in the Divadarreh region, Kordestan and near Borujerd, the Armand River in Chaharmahal va Bakhtiari Province (Raissy et al., 2010), Lorestan (records need verification by specimens), the Tedzhen River basin including the Jam and Kashaf rivers (Berg, 1949). This variable species is also recorded from the Dasht-e Kavir, Bejestan. Sirjan and  Namak Lake basins (Esmaeili et al., 2011?).

The subspecies Capoeta capoeta heratensis (the Transcaspian khramulya) is found in the Tedzhen or Harirud basin of Iran and eastwards including the Kashaf River (Abdoli, 2000). This subspecies is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Caspian Sea basin.

Zoogeography

Saadati (1977) suggests that this species entered the Dasht-e Kavir basin from either the Tigris River basin, the Hari River basin or the Caspian Sea basin. See also above under genus.

Habitat

The habitat of this species in the Surkhandar'ya of Uzbekistan is backwaters and channels with weak current and silt beds as well as reservoirs (Amanov, 1970). In Iran, it is one of two most abundant species in Caspian rivers along with Alburnoides bipunctatus (= eichwaldii) (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998). Ghasemi and Mustafayev (2008) found this species in the Aras River was the most dispersed and had the highest frequency (56.6%) of 17 species collected. It was found to be resistant to environmental changes, such as flooding, in the Madarsoo River in Golestan (Rezaei et al., 2008). Günther (1899) found that Capoeta capoeta placed in saline Lake Orumiyeh water died in 3.5 minutes.

Age and growth

Life span is over 8 years and catches in Uzbekistan are dominated by fish 3-4 or 4-5 years old. Growth is fastest in the first two years of life (Amanov, 1970). Life span in Azerbaijan is over 6 years (Abdurakhmanov, 1962), in northern Anatolia 6 years where vertebrae followed by scales were the best structures for aging (Polat and Işik, 1995), while in Georgia life span exceeds 9 years (Elanidze, 1983), and in Lake Gotchka, Armenia 10 years but only 4+ years in Lenkoran (Bănărescu in Bănărescu, 1999). Günther (1899) reports on a male fish from Ula in the Lake Orumiyeh basin which was only 12.5 cm long yet a sexually mature male, perhaps an instance of a dwarf form. Canbolat et al. (1999) found life span to be over 9 years in Çıldır Lake, Turkey for Capoeta capoeta capoeta. Fish aged 6 years dominated at 31.5% and 61.7% of the sample was female.

In Madarsoo Stream of Golestan National Park, this species had age groups 0-10 years and growth parameters were Lt = 229.67 mm and K = 0.54 for males, 327.95 mm and 0.18 for females (Koohestan Eskandari, 2003). Rezaei et al. (2007) also examined this fish population in the Madarsoo after two floods in 2001-2002. Growth parameters were L = 249 mm, K= 0.22 and t0 = -0.30 for males and  L = 306 mm, K= 0.21 and t0 = -0.38 for females. Length-weight relationships were W = -4.48 + 3.03TL for males and W = -4.59 + 3.0551TL in females, showing good feeding condition and positive isometric growth. Males were smaller than females as they matured earlier. Male to female ratio was 1.5:1, significantly different. Age range was 1+ to 5+ for males and 2+ to 8+ years for females. The dominant age was 2+ years.  Length was greatly decreased compared to previous studies and the population was younger, attributed to the floods. Shamekhi et al. (2012a) back-calculated lengths at previous ages for a population in the Dough Stream of the southeastern Caspian Sea basin; calculated lengths were smaller than observed lengths. Life span was 6 years. Shamekhi et al. (2012b) examined fish from five streams in the Gorgan River basin and found males to dominate (1.78:1), maximum relative abundance was 8.9-10.1cm for males and 7.7-8.9 for females, males were absent in the larger length groups, and there were various differences between streams and sexes for length groups and abundance, attributed to differences in food resources, growth rates, and selection favouring larger size in some streams.

Gholizadeh et al. (2009) studied a population in the Zarrin-Gol Stream in Golestan amd found age group 0+ was the most common at 59% and age groups 3+ and 4+ were the least common at 1%. Instantaneous growth of fish at age 3+ was much lower than younger age groups. The length-weight relationship was W = 0.00003xL2.822 and the von Bertalanffy equation was Lt = 223.8 (1-exp[0.185(t+1.8)]. Ranjbar et al. (2012) found fish in the Dough and Zarrin-gol streams to be 3-6 years old.

Maku Dam lake in West Azarbayjan has an estimated 9.4-10.7 tonnes of this species with a maximum sustainable yield of 4.5-4.8 t (Saiad Bourani and Ghaninejad, 2004). Average length of this population was 23.9 cm, weight was 1626.8 g and age was 2.6 years. Most fish were 3+ years old and 5+ fish were at a minimum. Infinite length and the growth coefficient were computed as 35.6 cm and 0.39 per year. Total mortality was 0.74, natural mortality 0.37 and fishing mortality 0.37. The Yasalegh Stream in the Gorgan River basin had a male to female ratio of 1:0.54, a maximum weight of 71.2 g for males and 119.4 g for females, and age range of 0-3 years, von Bertalanffy growth equations of Lt = 190(1-exp{-0.462 [t+1]} for males and Lt = 230(1-exp{-0.472 [t+0.742]} for females, and weight growth was isometric (b = 3.052 for males and 3.050 for females). Tilabad River fish had an age structure of 0-4 years and the Talar River 2-4 years, similar to Yasalegh Stream but differing from the Madarsoo Stream. The fish in the latter stream had better living conditions in a national park, no pollution, no fishing, no competition from exotic carps, no other human disturbances, no environmental stress and no food shortages. Patimar et al. (2009) found b values ranging from 2.647 (male at Chekchai) to 2.964 (females at Madarsu) indicating negative allometric growth for fish in the Gorgan River basin. They interpreted this variation to the species' response to different habitat conditions.

In the Tajan River fish showed negative allometric growth with W = 0.0138L2.9123 (Patimar et al., 2012).

Food

Food is mainly detritus and ooze, with some higher plants and small amounts of blue-green algae, which is digested in an intestine almost 7 times longer than the body (Amanov, 1970). Small benthic invertebrates may also be included, such as chironomids and molluscs (Bănărescu in Bănărescu, 1999). In Maku Dam lake, this species is a detritivore consuming Chrysophyta from the phytoplankton and Cyclotella a diatom, from the benthos as well as Chironomidae and Ephemeroptera (Valipour, 2004). In the Talar and Yasalegh rivers of the eastern Caspian Sea basin, 27 genera of phytoplankton were identified in the diet, with Chrysophyta being dominant, but with some differences between older and younger fish in the species consumed (Mostafavi and Abdoli, 2005).

Reproduction

Sexual maturity in Uzbekistan is at ages 2-4 years and lengths of 15-20 cm or in some populations at 10-14 cm. Some fish mature as dwarfs before age 1 and Berg (1948-1949) reports males 8.4 cm long of the subspecies gracilis can be mature. Spawning may take place at any time during the period from March to September (Berg,1948-1949) and is intermittent with the first spawning accounting for up to 85% of the eggs and the subsequent two spawnings for the remainder. The yellow eggs have a diameter up to 2.2 mm in the first spawning, up to 0.75 mm in the second and 0.65 mm in the third. Fecundity is up to 86,800 eggs. Eggs are laid at 50-60 cm on sand and stone beds and in water temperatures of 16-23°C (Amanov, 1970). Fecundity in the Kura River may reach 93,861 at 36-40 cm but this is for C. capoeta capoeta and fecundity for C. capoeta sevangi is less (Bănărescu in Bănărescu, 1999). Eggs are shed in running water and on lake shores, and eggs are covered by sand or small stones.

Rezaei et al. (2007) found no change in reproductive characteristics after floods in the Madarsoo Stream population. Mean fecundity was 3116 eggs and the maximum gonadosomatic index was in June. Ranjbar et al. (2012) found fish in the Dough and Zarrin-gol streams to have an average absolute fecundity of of 6030 and 5512 eggs respectively, the variation assumed to be different responses to different habitats. Absolute fecundity reached 9875 eggs and 12,107 eggs, relative fecundity 336 eggs/g and 252 eggs/g, with a maximum egg diameter of 1.03 mm and 1.18 mm.

Parasites and predators

Williams et al. (1980) report the helminths Khawia armeniaca (a cestode) and Acanthocephalorhynchoides cholodkowskyi (an acanthocephalan) from this species in the Zayandeh River at Esfahan. Molnár and Jalali (1992) record for this species the monogeneans Dactylogyrus chramulii, D. gracilis and D. lenkorani in the Safid Rud, D. chramulii and D. lenkorani in the Beshar River of the Persian Gulf drainage, D. gracilis and D. lenkorani in the Zayandeh Rud, D. lenkorani in the Tonekabon and Tajan rivers of the Caspian Sea drainage and the Kor River drainage of Fars, and D. pulcher from the Safid, Tajan, Tonekabon and "Ghasemlu" rivers of the Caspian Sea basin and the Jajrud of the Namak Lake basin. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species, the highest rate in 9 species examined. Malek (1993) and Malek and Mobedi (2001) report Clinostomum complanatum from this species in Mazandaran, the Shiroud. Up to 60 parasites per fish are recorded, with female fish having the highest infestation (the later study showing no difference between male and female fish), infestation decreasing with increase in body length, and parasites being concentrated in the gill cavity and pharynx.

Masoumian and Pazooki (1998) surveyed myxosporeans in this species in Gilan and Mazandaran provinces, finding Myxobolus musayevi and M. samgoricus. The crustacean parasite Tracheliastes polycolpus is reported from the fins of this species in the Mahabad Dam reservoir (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000). Jafari et al. (2001) isolated the acanthocephalan Dendronucleata dogieli from fish in the Zarrineh River, West Azerbaijan. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Myxobolus musayevi which is also recorded from this fish in the Tajan River in Mazandaran. Mokhayer et al. (2002) report Acanthocephalorhynchoides cholodkowskyi (Quadrigyridae) from the midgut and Tracheliastes polycolpus (Lernaeopodidae) on the fins of this fish in Golestan National Park, with more parasites on male fish and differences by season and station. Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Trichodina sp. and the monogenean trematode Dactylogyrus lenkorani. Mirhasheminasab and Pazooki (2003) list Ergasilus peregrinus, Tracheliastes polycolpus and Lernaea cyprinacea from this species in Mahabad Reservoir, the latter being the most dangerous parasite. Rohei Aminjan and Malek (2004) found 9 parasite species in fish from the Shiroud, namely the trematodes Clinostomum complanatum, Diplostomum spathaceum, Posthodiplostomum cuticola, Allocreadium sp., the monogeneans Dactylogyrus pulcher, D. lenkorani, Gyrodactylus mutabilitas and the nematodes Rhabdochona fortunatowi and Capillaria sp. Masoumian et al. (2005) recorded the protozoan parasites Ichthyophthirius multifilis, Trichodina perforata, Chilodonella, sp., Amphileptus branchiarum, Tetrahymena pyriformis, Apiosoma sp., and Vorticella sp. from this species in water bodies in West Azarbayjan. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus gracilis, D. charmulii, D. lenkorani and D. kendalanicus from this species in the Mahabad River of the Lake Orumiyeh basin, the latter species being a new record for Iran. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, namely Diplostomum spathaceum, Ligula intestinalis, Digrama sp., Rhabdochona hellichi, Argulus foliaceus, Allocreadium isoporum, Lamprolegna compacta, Myxobolus cristatus and M. musajevi. Pazooki et al. (2005) record Tracheliastes longicollis, Lamprolegna compacta, Neoechinorhynchus rutili, Capillaria sp., Myxobolus musajevi, M. cristatus, Trichodina perforata, Chilodonella piscicola, Ichthyophthirius multifilis and Ichthyobodo necatrix from this species in waterbodies of Zanjan Province. Pazooki et al. (2006) record the monogeneans Dactylogyrus chramuli, D. gracilis, D. lamellatus, D. lenkorani, D. pulcher and Gyrodactylus sp. from this fish in Zanjan Province. Masoumian et al. (2007) record the myxosporean parasite Myxobolus musajevi from this species in the Zayandeh River. Miar et al. (2008) examined fish in Valasht Lake and the Chalus River, Mazandaran and found the metazoan Myxobolus saidovi. Maleki and Malek (2007) examined fish from the Shirud in the Caspian Sea basin and recorded the digeneans Posthodiplostomum cuticola, Diplostomum spathaceum, Clinostomum complanatum and Allocreadium sp. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus foliaceus, Ergasilus sp., Lamproglena compacta, Lernaea sp., Tracheliastes longicollis and Tracheliastes polycolpus on this species. Raissy et al. (2010) found ichthyophthiriasis (infection with Ichthyophthirius multifilis - ich or white spot disease), which cause epizootics in wild and cultured fishes, in fish from the Armand River in Chaharmahal va Bakhtiari Province.

Economic importan

The subspecies Capoeta capoeta heratensis is a food fish in Uzbekistan (Amanov, 1970) and C. capoeta sevangi and C. capoeta capoeta are commercially important in Lake Gotchka, Armenia and eastern Georgia and Azerbaijan respectively (Bănărescu in Bănărescu, 1999). It is also used in sport fishing in Iran (Samaee et al., 2006). Banagar et al. (20110 report concentrations of cadmium in liver and muscle tissue for fish from the Tajan River higher than standard limits. Shiri et al. (2009) and Ramin et al. (2012) report a case of ichthyotoxism after eating fried eggs of this species from Marmisho Lake west of Orumiyeh. Nausea resulted after one minute, and the victim was hospitalised with severe chest pains. No vomiting occurred as this was the only food eaten and symptoms appeared rapidly. Raw consumption should be avoided and even cooked fish or inadequately cleaned fish can be dangerous.

Conservation

Kiabi et al. (1999) consider this species to be of least concern in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin.

Further work

The relationships of the various subspecies need study to determine if they are in fact good species.

Sources

Type material: See above, Capoeta gibbosa (ZISP 11104).

Iranian material: CMNFI 1970-0512, 3, mm standard length, Gilan, Shalman River (37º08'N, 50º15'E); CMNFI 1970-0514, 17, ? mm standard length, Gilan, Shafa River estuary (37º55'N, 49º09'E); CMNFI 1970-0516, 6, ? mm standard length, Gilan, Lemir River (38º14'N, 48º52'30"E); CMNFI 1970-0519, 2, ? mm standard length, Gilan, Chelvand River (ca. 38º18'N, ca. 48º52'E); CMNFI 1970-0520, 2 ?7 on data sheet, 94.0-100.9 mm standard length, Gilan, Astara River (ca. 38º25'N, ca. 48º52'E); CMNFI 1970-0521, 3, ? mm standard length, Gilan, Safid River near Lulaman (no other locality data); CMNFI 1970-0522, 10, ? mm standard length, Gilan, Safid River at Astaneh bridge (37º16'30"N, 49º56'E); CMNFI 1970-0525, 5, 92.8-146.1 mm standard length, Gilan, Safid River near Mohsenabad (ca. 37º22'N, ca. 49º57'E); ?see data sheets 78, 22.1-160.4 CMNFI 1970-0526, 8, ? mm standard length, Gilan, Safid River 6 km below Astaneh bridge (37º19'N, 49º57'30"E); CMNFI 1970-0531, 7, 60.2-84.6 mm standard length, Mazandaran, Larim River (36º46'N, 52º58'E); CMNFI 1970-0536, 2, ?5 on data sheets 101.4-125.4 mm standard length, Gilan, Siah River estuary (36º53'N, 49º32'E); CMNFI 1970-0538, 1, ?5 on data sheet 95.4 mm standard length, Gilan, Qezel Owzan River near Manjil (36º44'N, 49º24'E); CMNFI 1970-0557, 3, ? mm standard length, Azarbayjan-e Bakhtari, Shaher Chay (no other locality data); CMNFI 1970-0559, 6, 83.9-125.4 mm standard length, Azarbayjan-e Bakhtari, Baranduz Chay (ca. 37º25'N, ca. 45º10'E); CMNFI 1970-0568, 9, ? mm standard length, Gilan, Caspian Sea at Kazian beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0577, , mm standard length, Gilan, Caspian Sea at Astara (ca. 38º26'N, ca. 48º53'E); CMNFI 1970-0583, 8, 34.1-93.9 mm standard length, Gilan, Nahang Roga (37º28'N, 49º28'E); CMNFI 1979-0589, , mm standard length, Gilan, Safid River opposite Kisom (37º12'N, 49º54'E); CMNFI 1979-0242, 27, 25.6-107.0 mm standard length, Fars, river at Izadkhvast (31º31'N, 52º07'E); check ID? CMNFI 1979-0249, 33, 66.4-114.2 mm standard length, Esfahan, stream at Dizaj (31º55'N, 51º30'E); check ID? CMNFI 1979-0429, 1, ? mm standard length, Mazandaran, Chalus River (36º34'N, 51º23'E); CMNFI 1979-0432, 1, ? mm standard length, Mazandaran, Sardab River branch (36º41'N, 51º22'E); CMNFI 1979-0434, 1, ? mm standard length, Mazandaran, Shir River (36º51'N, 50º49'E); CMNFI 1979-0435, 1, ? mm standard length, Gilan, stream 10 km west of Ramsar (36º57'N, 50º37'E); CMNFI 1979-0433, 1, 115.2 mm standard length, Mazandaran, stream 18 km west of Chalus (36º42'N, 51º15'E); CMNFI 1979-0438, 2, 142.4-144.8 mm standard length, Gilan, Gholab Ghir River (37º27'N, 49º37'E); CMNFI 1979-0441, 1, 121.9 mm standard length, Gilan, river 14 km south of Hashtpar (37º42'N, 48º58'E); CMNFI 1979-0443, 1, ? mm standard length, Gilan, river 34 km west of Hashtpar (38º06'N, 48º53'E); CMNFI 1979-0444, 1, ? mm standard length, Gilan, Chubar River (38º11'N, 48º52'30"E); CMNFI 1979-0446, 1, ? mm standard length, Gilan, Astara River (38º26'30"N, 48º51'E); CMNFI 1979-0449, 2, ? mm standard length, Azarbayjan-e Khavari, river 18 km from Khalkhal (ca. 37º42'N, ca. 48º27'E); CMNFI 1979-0451, 30, 35.8-97.3 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River (ca. 37º30'N, ca. 47º57'E); CMNFI 1979-0452, 1, 79.7 mm standard length, Azarbayjan-e Khavari, Qezel Owzan River 6 km from Mianeh (37º23'N, 47º45'E); CMNFI 1979-0453, 24, 36.1-111.1 mm standard length, Zanjan, Zanjan River (37º06'N, 47º56'E); CMNFI 1979-0469, 2, 56.6-76.2 mm standard length, Mazandaran, river 36 km west of Alamdeh (36º37'30"N, 51º35'E); CMNFI 1979-0474, 1, ? mm standard length, Mazandaran, Tajan River (36º34'N, 53º05'E); CMNFI 1979-0475, 1, 86.4 mm standard length, Mazandaran, stream on road to Bandar-e Shah (36º46'N, 54º00'E); CMNFI 1979-0480, 2, ? mm standard length, Mazandaran, Gorgan River at Gonbad-e Kavus (37º15'30'N, 55º09'E); CMNFI 1979-0481, 3, 101.9-188.0 mm standard length, Mazandaran, stream 3 km west of Ghalahleekesh (37º18'30"N, 55º31'E); CMNFI 1979-0482, 2, ? mm standard length, Mazandaran, river 2km west of Ghalahleekesh (37º19'30'N, 55º31'E); CMNFI 1979-0483, 4, 121.6-160.5 mm standard length, Mazandaran, river 28 km west of Dasht (37º23'30"N, 55º51'30"E); CMNFI 1979-0485, 3, 71.2-99.1 mm standard length, Khorasan, stream 28 km west of Bojnurd (37º33'N, 57º04'E); CMNFI 1979-0486, 66, 17.5-97.8 mm standard length, Mazandaran, stream in Atrak River draiange (37º44'N, 56º18'E); CMNFI 1979-0487, 20, ? mm standard length, Mazandaran, spring 2 km from Maraveh Tappeh (37º54'N, 55º58'E); CMNFI 1979-0488, 9, 29.7-140.4 mm standard length, Mazandaran, Atrak River at Maraveh Tappeh (37º55'N, 55º57'30"E); CMNFI 1979-0489, 78, ? mm standard length, Mazandaran, stream 13 km from Maraveh Tappeh (37º50'N, 55º53'E); CMNFI 1979-0490, 14, 21.0-108.4 mm standard length, Mazandaran, stream in Gorgan River drainage (ca. 37º39'N, ca. 55º42'E); CMNFI 1979-0491 2, ? mm standard length, Mazandaran, Gorgan River northeast of Kalaleh (ca. 37º33'N, ca. 55º44'E); CMNFI 1979-0492, 25? check jar, 9.3-183.4 mm standard length, Mazandaran, river in Gorgan River drainage (37º05'N, 55º15'E); CMNFI 1979-0695, 13, ? mm standard length, Gilan, Safid River at Manjil Bridge (36º46'N, 49º24'E); CMNFI 1997-0003, , mm standard length, (); CMNFI 1980-0116, 1, ? mm standard length, Gilan, Safid River at Astaneh (37º16'30"N, 49º56'E); CMNFI 1980-0120, , mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1980-0121, , mm standard length, Gilan, Shafa River estuary (37º35'N, 49º09'E); CMNFI 1980-0123, , mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E); CMNFI 1980-0141, , mm standard length, Gilan, Lisar River estuary (37º59'N, 48º56'E); CMNFI 1991-0163, , mm standard length, Mazandaran, Ramian River (36º58'N, 55º07'E); CMNFI 1993-0138, 1, mm standard length, Khorasan, Bazangan Lake (36º18'N, 60º27'E); CMNFI 2007-0014, 4, 39.4-99.1 mm standard length, Khorasan, pool in Kuh-e Sang Park, Mashhad (ca. 36º18'N, ca. 59º36'E); CMNFI 2007-0086, 6, ? mm standard length, Azarbayjan-e Khavari, Qareh Su basin near Nir (ca. 38º02'N, ca. 48º00'E); CMNFI 2007-0087, 1, ? mm standard length, Azarbayjan-e Khavari, Qareh Su north of Ardebil (38º22'N, 48º19'E); CMNFI 2007-0088, 5, ? mm standard length, Azarbayjan-e Khavari, Qareh Su east of Lari (38º30'N, 48º03'E); CMNFI 2007-0089, 4, ? mm standard length, Azarbayjan-e Khavari, Ahar Chay at Ahar (38º28'N, 47º03'E); CMNFI 2007-0093, 13, ? mm standard length, Azarbayjan-e Bakhtari, Qotur River south of Khvoy (38º30'N, 44º58'E); CMNFI 2007-0094, 6, ? mm standard length, Azarbayjan-e Bakhtari, Nazlu River north of Reza'iyeh (ca. 37º42'N, ca. 45º04'E); checkID? CMNFI 2007-0095, 2, ? mm standard length, Azarbayjan-e Bakhtari, Shahr Chay southwest of Reza'iyeh (ca. 37º27'N, ca. 44º56'E); checkID? CMNFI 2007-0096, 5, ? mm standard length, Azarbayjan-e Bakhtari, Qasemul River in Baranduz Chay basin (ca. 37º25'N, ca. 45º10'E); checkID? CMNFI 2007-0098, 2, ? mm standard length, Azarbayjan-e Bakhtari, river south of Mahabad (ca. 36º42'N, ca. 45º41'E); CMNFI 2007-0099, 1, ? mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay west of Mahabad (ca. 36º35'N, ca. 45º25'E); checkID? CMNFI 2007-0101, 1, ? mm standard length, Azarbayjan-e Bakhtari, Tata'u River south of Miandow Ab (ca. 36º54'N, ca. 46º07'E); CMNFI 2007-0102, 4, ? mm standard length, Azarbayjan-e Bakhtari, Zarineh River near Miandow Ab (ca. 37º00'N, ca. 46º07'E); CMNFI 2007-0103, 9, ? mm standard length, Kordestan, Zarineh River basin north of Saqqez (ca. 36º18'N, ca. 46º16'E); CMNFI 2007-0104, 4, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º12'N, ca. 46º18'E); CMNFI 2007-0105, 7, ? mm standard length, Kordestan, Zarineh River basin south of Saqqez (ca. 36º06'N, ca. 46º20'E); CMNFI 2007-0106, 9, ? mm standard length, Kordestan, Qezel Owzan River basin near Divandarreh (ca. 35º52'N, ca. 47º05'E); CMNFI 2007-0107, 10, ? mm standard length, Kordestan, Qezel Owzan River basin near Bijar (ca. 35º54'N, ca. 47º20'E); ZSM 24500, 6, 24.2-31.0 mm standard length, Khorasan, stream near Bojnurd (no other locality data); uncatalogued, 2, 75.8-85.5 mm standard length, Khorasan, Hari River at Sarakhs (36º32'N, 61º11'E).

Capoeta damascina
(Valenciennes, 1842)

Zayandeh River, February 2008, courtesy of K. Abbasi
Zayandeh River, February 2008, courtesy of K. Abbasi

Common names

sardeh (= cold one, probably zardeh = yellow one, is more correct and appropriate based on yellow-tinged flank) or سياه ماهي (= siah mahi, meaning black fish) in the Caspian basin; siah mahi damascina; tu'ini (meaning unknown) or gel cheragh (= mud-eater, mud-grazer) or tu'ini gelkhorak (= mud-eater, mud-grazer) in Khuzestan; qezel ala (red spots) in Chahar Mahall but mistakenly.

[twena, toyoueni or toueni, bertin or bartin, tin, zardah masih, tela shami, tela Damascus; kollur, kellur, kollur hadjiari (= the pilgrim or migrating kollur), kellur dischileki (= the strawberry-coloured kollur), kollur achmar (= the red kollur) and kollur aschkar (= the brown kollur), all at Aleppo; all in Arabic; Mesopotamian barb].

Systematics

Gobio damascinus was described from the "fleuve de Damas" (= river of Damascus, Syria).

Synonyms are Scaphiodon capoeta Heckel, 1843 (non sensu Güldenstädt, 1773) described from "Aleppo", Scaphiodon fratercula Heckel, 1843 described from "Gewässern von Damascus", possibly Scaphiodon Umbla Heckel, 1843 described from the "Tigris bei Mossul", Scaphiodon socialis Heckel, 1843 described from "Um Damascus" (= around Damascus) (Heckel, 1843b) and later more completely from the "Orontes" (Heckel, 1847a) (placed in Scaphiodon Capoëta of Heckel by Steindachner (1864)), Scaphiodon peregrinorum Heckel, 1843 described from "Aleppo" and later from "Fluss Kueik bei Aleppo", Chondrostoma syriacum Valenciennes, 1844 from Abraham's River at the foot of Mount Sinai, Egypt (the correct locality is probably in the Jordan River basin (Coad and Krupp, 1994)), Scaphiodon Amir Heckel, 1847 described from the "Araxes" (= Kor River, Fars), Scaphiodon niger Heckel, 1847 described from the "Araxes oder Benth-Amir" (= Kor River, also known as the Bandamir River), Scaphiodon Saadii Heckel, 1847 described from the "Quellen des Saadi" (Sa`di at 29°37'N, 52°35'E, now within the city of Shiraz) and the "Nähe von Persepolis" (= probably the Pulvar (= Sivan) River near Persepolis, Fars), Scaphiodon chebisiensis Keyserling, 1861 from "Wasserleitung in Chebis" (= canal in Chebis, probably Khabis or Shahdad at 30°25'N, 57°42'E in Kerman), Scaphiodon rostratus Keyserling, 1861 from "Wasserleitungun in der Umgegend von Jezd. Das abgebildete Exemplar stammte aus Meibut" (= canals in the vicinity of Yazd. The specimen drawn originated from Meibut, probably Meybod at 32°14'N, 54°01'E), Barbus belayewi Menon, 1960 (Menon and Yazdani (1968) date this species as 1960, presumably the 1956 edition of the journal was delayed) from the "Tigris, Baghdad, Iraq", and Capoeta capoeta intermedia Bianco and Banarescu, 1982 (non Capoeta intermedia Temminck and Schlegel, 1846 = Acheilognathus lanceolata (Temminck and Schlegel, 1846) (see Boeseman, 1947)) described from the "Mand River near Akbar, southern Iran".

The synonymy of Barbus belayewi is suggested by F. Krupp (in litt., 1986) and W. Rainboth (pers. comm., 1986). The synonymy of S. fratercula is pointed out by Berg (1949) since the species was founded on low lateral line scale counts, a variable character in C. damascina, and on a larger orbit but Heckel's comparison was between fish of greatly differing size and no allowance was made for allometry.

Karaman (1969) places damascina in Capoeta capoeta as a subspecies and umbla as another subspecies. Berg (1949) and Saadati (1977) recognise umbla as a distinct species. The latter is distinguished from the former by a higher scale count (87-99), higher dorsal fin branched rays (9-10), longer dorsal fin, longer caudal fin (shorter than or equal to head length in C. damascina), a markedly transverse mouth, and a weaker dorsal fin spine. Saadati (1977) considers fratercula to be a distinct species from the Tigris and Mand rivers in Iran based on scale count (58-66), more gill rakers (20-22), and a more serrated dorsal fin spine; or a subspecies of Capoeta capoeta based on a close similarity in scale counts, average number of gill rakers, and the dorsal fin origin being anterior to that of the pelvic fins. He also considers that Scaphiodon niger from the Kor River of Fars is possibly a synonym of fratercula. Krupp (1985c) considers the synonymy of C. damascina and C. capoeta as extremely doubtful after examining topotypic material.

Bianco and Banarescu (1982) recognise Capoeta saadi as a distinct species based on an arched mouth rather than transverse as in most subspecies of Capoeta capoeta, with a lightly developed horny cover on the lip, a feebly ossified dorsal fin spine, 13-17 gill rakers, modally 8 dorsal fin branched rays, 53-76 lateral line scales and 24-28 scales around the caudal peduncle. However they do point out the extreme variability in scale counts, for example, from fish taken in the same locality and even between opposite sides of the same fish (5 more scales on one side than the other!). Designation of subspecies on such variable characters is difficult and would require very large series and multivariate analysis techniques. Bianco and Banarescu (1982) regard C. c. intermedia as intermediate between C. c. umbla and their C. c. macrolepis on the basis of scale counts, gill raker counts, smaller transverse mouth than in umbla and a rather light colouration.

Capoeta damascina, with a wide distribution and wide variation in morphology, must be regarded as a species complex until detailed analyses can be carried out. Final resolution of the species composition of this complex may well require extensive material for molecular analyses, as well as re-examination of types and topotypic material over the whole range of the taxon. Samaee and Patzner (2011) examined fish from 6 river systems in Iran morphometrically and were able to distinguish distinct groups. However, as they point out, much more work needs to be done to determine if this variation is genetic differentiation or phenotypic plasticity, or a combination of the two.

The syntypes of Gobio damascinus are in the Muséum national d'Histoire naturelle, Paris (MNHN 4494, 2 specimens, 169-179 mm standard length, Damascus, Syria, Bové, MNHN 3948, 1, 289 mm standard length, Nahr Barada, Syria and MNHN A.3947, 1, 169 mm standard length, Syria) (Krupp, 1985c). Bertin and Estève (1948) give 200-210 mm total length for MNHN 4494 and 330-390 mm total length for MNHN 3947, 3948 and A.789. Eschmeyer et al. (1996) list MNHN 4494 as the lectotype (as designated by Krupp and Schneider (1989) although this collection comprises two fish) with MNHN 3947 (1, dry) and MNHN 3948 (1, dry) and possibly MNHN A.789 (1) as paralectotypes. The latter is listed as a syntype in Bertin and Estève (1948) although the localities listed in this article "Fl. Jourdain, à Damas (Syrie)" is obviously an error on geographical grounds.

Syntypes of Scaphiodon capoeta are in the Naturhistorisches Museum Wien under NMW 51650 (1 fish), NMW 51831 (1), and NMW 55845-55846 (2). Heckel (1843) lists 2 specimens in his description.

The holotype of Chondrostoma syriacum is in the Muséum national d'Histoire naturelle, Paris under MNHN 1945 (Eschmeyer et al., 1996).

The holotype of Capoeta capoeta intermedia is in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (IZA 7892) and is 92.5 mm standard length, collected by P. Bianco and S Zerunian, 27/5/1976. There are 62 paratypes (IZA 7893) from the same collection as the holotype measuring 36-87 mm standard length and 13 paratypes uncatalogued in the Institutul de Stiinte Biologice, Bucuresti, Romania (ISBB) measuring 68-86 mm standard length (Bianco and Banarescu, 1982). Another paratype under IZA 7894 measures 105.5 mm standard length was examined by me. A paratype of Capoeta capoeta intermedia from the Mand River in Fars is in the Zoologischen Instituts und Zoologischen Museums der Universität Hamburg (ZMH 6090, 83.2 mm standard length) (Wilkens and Dohse, 1993; examined by me), one paratype from the Mand is in the California Academy of Sciences, San Francisco (CAS 48113), one paratype from the Mand is in the United States National Museum, Washington (USNM 227935), and 6 paratypes are in the Canadian Museum of Nature, Ottawa under CMNFI 1982-0367 (formerly IZA 7893).

The holotype of Scaphiodon fratercula was taken from "Gewässern von Damascus", the syntypes of Scaphiodon umbla from the "Tigris bei Mossul", the types of Scaphiodon socialis from "Um Damascus" (but listed as "Orontes" in the catalogue in Vienna, possibly in confusion as this part of the catalogue has been overwritten), and the types of Scaphiodon peregrinorum from "Um Aleppo" according to Heckel (1843b) and "Fluss Kueik bei Aleppo" according to Heckel (1847a).

Two syntypes of Scaphiodon niger are in the Naturhistorisches Museum Wien under NMW 51655 with standard lengths of 140.4 and 188.5 mm (another syntype is under NMW 51654 (232.7 mm), and a fourth under NMW 51656 as seen by me; all 4 are listed as syntypes in the 1997 Vienna card index). Eight syntypes of Scaphiodon amir are under NMW 61472 and measure 42-59 mm standard length and there are also 6 fish under NMW 46081 (138.1-282.3 mm standard length); however the card index in 1997 lists only NMW 46081 (6) and 16508 (1, dried). Fifteen syntypes in the catalogue (18 seen by me and in the Vienna card index in 1997) of Scaphiodon saadii from Sa`di are under NMW 51666 (Eschmeyer et al. (1996) have 52666, apparently in error) and measure 58-123 mm standard length (18.3-123.8 mm standard length when measured by me) with a further 4 syntypes from Persepolis under NMW 55900 measuring 84-114 mm standard length (Kähsbauer, 1964; not in the 1997 card index). There is also 1 syntype (RMNH 3166) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW (Eschmeyer et al., 1996).

The catalogue in Vienna lists no fish opposite the name S. niger, 6 and 2 fish in one column and 5 in the adjacent column for Scaphiodon amir (cf. above), 10 fish in one column and 10 in the adjacent column for S. saadii (cf. above).

A dried syntype of C. umbla is in the Senckenberg Museum Frankfurt (SMF 6777, formerly NMW) (F. Krupp, pers. comm., 1985; ca. 262.3 m standard length), 2 syntypes are in Naturhistorisches Museum Wien (NMW 55932-55933) and another syntype is under NMW 55934. Eschmeyer et al. (1996) also lists NMW 79373-74, both dried. The catalogue in Vienna lists 2 fish in spirits and 2 fish stuffed and the card index in 1997 lists as syntypes NMW 55932-33 and 79373-74 (dried).

Eschmeyer et al. (1996) note that there are no types of Scaphiodon fratercula in the Naturhistorisches Museum Wien.

Two fish are labelled as syntypes of Scaphiodon socialis in the Naturhistorisches Museum Wien (NMW 55855) which agrees with Heckel's text although the catalogue lists only 1 specimen. Eschmeyer et al. (1996) state that there are no types at NMW presumably after Krupp and Schneider (1989) who state that NMW 55670 (1 fish), 55843 (2) and 55855 (2) are not types.

The types of Scaphiodon peregrinorum number 6 according to the catalogue in the Naturhistorisches Museum Wien and may comprise all or part of NMW 51658 (1), NMW 51659 (1), NMW 51660 (1), NMW 51661 (1), NMW 51662 (1), NMW 51663 (1) NMW 51664 (3), and NMW 51665 (1), all labelled as from "Kueik" and possibly RMNH 2681 (3) in the Rijksmuseum van Natuurlijke Historie, Leiden from NMW (Eschmeyer et al., 1996).

The types of Barbus belayewi are in the Zoological Survey of India, Calcutta, the holotype being ZSI F1046/2 and a paratype ZSI F1047/2 (Menon, 1960; Menon and Yazdani, 1968).

Types of Scaphiodon chebisiensis and Scaphiodon rostratus were not kept.? phrasing

Key characters

The mode of 9 dorsal fin rays, small scales, and the presence of large black blotches often distinguish this species from other Capoeta in Iran.

Morphology

Dorsal fin with 3-5 unbranched rays and 8-10 branched rays (Krupp (1985c) gives frequency distributions for his material from Turkey, Syria, Lebanon, Israel and Jordan as 8(52), 9(144) and 10(4)), anal fin with 3 unbranched and 5-6 branched rays (5(179), 6(21) after Krupp (1985c)), pectoral fin branched rays 15-20, and pelvic fin branched rays 8-10. Lateral line scales 60-99. Esmaeili et al. (2007) detail scale structure of fish from southwest Iran using scanning electron microscopy. Gill rakers 17-25 (Saadati (1977) gives 9-21!; Krupp (1985c) 12-18 for the lower arm of the arch, Berg (1949) up to 23 on the lower arm). Pharyngeal teeth 2,3,4-4,3,2, often 2,3,5-5,3,2, with spoon-shaped crowns. The mouth is usually horseshoe-shaped, seldom transverse. The last unbranched dorsal fin ray is moderate to strong with denticles along two-thirds of its length. Heckel (1847b) distinguished his Scaphiodon amir and S. niger by the dorsal fin denticles being horizontal or perpendicular to the spine, not hooked downward as in related species. Berg (1949) did not attach any significance to this character, finding it in small fish from the Sarhadd of Baluchestan and from Jordan.

The karyotype for fish in the Tigris River basin of Turkey identified as Capoeta capoeta umbla is 2n=150, possibly hexaploid (Kılıç Demirok and Ünlü, 2001) and of fish identified as C. damascina from the Wadi Karak, Jordan 2n=148-150, indicating a hexaploid species (Gorshkova et al., 2002).

Body form is highly variable as are scale counts between populations and even within populations when large series are examined (Krupp, 1985c). Subspecific designations can only be valid if very large series from the whole range of the species are compared.

Meristic values for Iranian specimens are :

IZA7892, 7894 (2 fish) D8, A5, P15-15, V7, ll 62-64, gr 24

Sexual dimorphism

Males develop breeding tubercles around the snout and the posterior body on both sides of the lateral line (Khalaf, 1987).

Colour

The back is dark brown or brownish to olive or blue-grey, the flanks silvery with some yellowish tinges, sometimes golden, or yellow-brown or reddish-brown above the lateral line, silvery below. The belly is white to yellowish. Cheeks are golden. Dark brown or black spots numbering up to 20 may be scattered irregularly on the flanks. Fins are reddish-brown, yellowish or grey and may be hyaline. The caudal and pectoral fins may be very dark compared to other fins. The pectoral and pelvic fins may a light pink tinge. The cartilaginous edge to the lower jaw is bright yellow to red-yellow. The peritoneum is black. Some fish may be very black with only the underside of the head and belly yellowish-white (specimens described by Heckel (1847b) as Scaphiodon niger; however since these fish "decompose quickly in the commonly used ethyl alcohol concentrations", they may have been poorly preserved and the black colouration resulted from partial decomposition).

Size

Attains 35.3 cm standard length, about 45.0 cm total length and 0.5 kg.

Distribution

Found from Turkey, Syria, Lebanon and Israel to Iran. In Iran, it is reported from the Tigris River including the the Regab River in Kurdistan and the Selakhor River near Borujerd, the Jarrahi, Marun, Karun, Kuhrang, Bazoft and Khersan rivers, throughout the Dez and Karkheh basins to their uppermost reaches, Gheshlagh Dam Lake, Sanandaj; Lake Zarivar; Armand River in Chaharmahal va Bakhtiari Province, Esfahan including the Dopolan and Zayandeh rivers, Dasht-e Kavir including the Jajarm and upper Kal Shur rivers, Namak Lake including the Karaj, Shur, Abhar, Qareh Su and Qom rivers; Kor River, Lake Maharlu, Gulf including the Zohreh River and its Kheirabad tributary and the Mand River near Akbar and its Shur (Dasht-e Palang) River tributary, Kerman-Na'in, Dasht-e Lut, Sirjan, Hormuz including the middle to upper Hasan Langi, Kul and its Shur River tributary, and Hamun-e Jaz Murian basins (Lovett, 1873; Nikol'skii, 1899; Berg, 1949; Kähsbauer, 1964; Spillman, 1972; Armantrout, 1980; Rainboth, 1981; Bianco and Banarescu, 1982; Abdoli, 2000; Ghorbani Chafi, 2000; R. Mehrani, pers. comm., 2000; Jalali et al., 2005; Raissy et al., 2010; Esmaeili et al., 2011?; Bahrami Kamangar et al., 2012).

Zoogeography

Its relationships with other Capoeta species is generally unclear, as is the status of isolated populations some of which have been named. The larger zoogeographical relationships of this and other Capoeta species remain uncertain. See also above under genus.

Habitat

Unknown in detail.

Age and growth

All males are mature at 18 cm and all females at 20 cm in Khalaf's (1987) study in the Lebanon. In Lake Kinneret, Israel, Stoumboudi et al. (1993) found that fish longer than 25 cm have developed gonads, occasionally males mature between 16 and 25 cm as did females between 20 and 25 cm. Khalaf et al. (2002) found 6 age classes (1+ to 6+) in the Nahr el Khalb, a Lebanese stream. Maximal growth was in July and August and minimal growth between December and February.

Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 40 Iranian fish measuring 5.23-19.87 cm standard length. The a-value was 0.0282 and the b-value 2.890 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Asadollah et al. (2011) found fish up to 10+ years in the Zayandeh River, the oldest fish being female, and the most frequent age classes being 3+ for males and 4+ for females.

Food

Khalaf (1985) and Spataru and Gophen (1986) examined the food of this species in Lebanon and Israel respectively. Benthic diatoms and filamentous algae are the main foods. Some other algal species and some zoobenthic organisms are present along with large quantities of mud. The species is classified as a phytobenthophagous fish, one that takes its food from bottom sediments. Leaf remains have also been found in gut contents. Abdoli (2000) lists variety of insects: Chironomidae, Formicidae, Epididae, Empididae, Tipulidae, Tabanidae, Simuliidae, Hydroptilidae, Grouvellinus, Elmis, Hydropsyche, Heptagenia, Baetis and hydracariens.

Reproduction

Al-Rudainy (2008) gives sexual maturity at 2-3 years in Iraq with spawning in May, absolute fecundity up to 53,000 eggs and relative fecundity up to 7300 eggs/g body weight, and average egg diameter 1.48 mm.

Khalaf (1987) examined the reproductive cycle in this species for Lebanese waters. Spawning begins in May and ends in July. Eggs number up to 5,138 and egg diameters are up to 2.2 mm. In marked contrast, Stoumboudi et al. (1993) found that gonad weights are greatest in January in Lake Kinneret, Israel, 4 months earlier. This may be evidence of different temperature regimes or populational variation. Fishelson et al. (1996) confirm that this species migrates in winter, December to February in the upper Jordan River of Israel, the process being initiated by rainfall and flooding and a decrease in temperature to 16-18°C. The gonadosomatic index is highest in February and the final months of reproduction are March to May. Lake dwelling fish aggregate and swim up streams as far as 25 km and altitudes of 400-900 m, fattening and ripening at the spawning site. They can jump rapids on this migration (and in Iran large fish cornered in small streams will jump over seine nets!). The females excavate a shallow nest in which to deposit adhesive eggs, up to 4.5 mm in diameter. Dozens of nests are found close together and sand and gravel stirred up by the excavation covers adjacent nests. After spawning the adults return downstream to the stream mouth and lake.

Asadollah et al. (2011) examined reproduction in fish from the Zayandeh River. The sex ratio was 1:1.57 (male:female). Males matured at 2+ years and females between ages 4 to 6 years with 100% maturity at age 7+ years. Gonadosmatic indices indicate reproduction in May and June at 12.6-16.6°C and it is group-synchronous. Absolute fecundity reached 72,645 eggs, with 4+ females averaging 9446 eggs and 8+ females 54,503 eggs. Mean egg diameter was highest in May at 1.81 mm.

Parasites and predators

Dollfus (1970) describes a new cestode Coelobothrium monodi from this species at "Nasratabad", possibly from the Dasht-e Lut basin. Jalali et al. (1995) describe two new species of monogeneans, Dactylogyrus rohdeianus and D. capoetae, from fish caught in the "Chaghalnandi" River, a Karkheh River tributary north of Ahvaz. González-Solís et al. (1997) report the nematodes Rhabdochona denudata and Rhabdochona fortunatowi from this species in the Mand River, Fars. O. M. Amin (pers. comm., 1998) has identified the acanthocephalan Acanthocephalorhynchoides cholodkowskyi from specimens collected in the Mand River west of Shiraz, Fars. Jalali et al. (2002) and Jalali and Barzegar (2006) record Trichodina pediculus, Dogielius molnari, Gyrodactylus sp., Dactylogyrus carassobabrbi and D. lenkorani from this species in Lake Zarivar. Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Dactylogyrus lenkorani, Gyrodactylus pulcher, Dactyolgyrus sp., Allocreadium isoporum and Myxobolus molnari. Mehdipoor et al. (2004) record the monogeneans Dactylogyrus lenkorani and D. pulcher in Zayandeh River fish. Barzegar and Jalali (2006) report a parasite in this species from Kaftar Lake as Dactylogyrus lenkorani. Masoumian et al. (2007) record the myxosporean parasites Myxobolus samgoricus and M. varicorhini from this species in the Zayandeh River. Barzegar et al. (2008) record the digenean eye parasite Diplostomum spathaceum from this fish. Nazari Chamak et al. (2010) found the following myxozoan parasites in the genus Myxobolus: buckei, cristatus, karelicus, musajevi, samgoricus, suturalis and varicorhini in fish from the Halil River, Kerman. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea cyprinacea on this species. Jalali Jafari and Miar (2011) summarise the metazoan parasite community in this species in the Mesopotamian region, identifying 54 species. Raissy et al. (2010) found ichthyophthiriasis (infection with Ichthyophthirius multifilis - ich or white spot disease), which cause epizootics in wild and cultured fishes, in fish from the Armand River in Chaharmahal va Bakhtiari Province.

Economic importance

This species is of no economic importance in Iran although in Israel annual catches in Lake Kinneret have been as high as 29 tonnes or 7% of the total fishery (Spataru and Gophen, 1986). Heckel (1847b) reports that this species was "greatly appreciated as food fish by the local people" in the Kor River basin, Fars (as his Scaphiodon amir). Samaee and Patzner (2011) mention that it is fished recreationally in Iran.

The eggs are reputedly poisonous and this is said to account for the low population of introduced Oncorhynchus mykiss in Gahaar Lake, Lorestan (R. Mehrani, pers. comm., 2000).

Conservation

A widely distributed species, probably not in need of conservation. Endangered in Turkey (Fricke et al., 2007). Bahrami Kanagar et al. (2012) provided baseline haematological and biochemical indices for this species in Gheshlagh Dam Lake, Sanandaj that could be used in health monitoring.

Further work

The relationships of isolated and named taxa under this species, and the relationships of this species to other Capoeta, would benefit from molecular analyses.

Sources

Type material: See above, Capoeta capoeta intermedia (IZA 7892, 7894, CMNFI 1982-0367 (formerly IZA 7893), ZMH 6090) P. G. Bianco. ? check this last speciemne; Scaphiodon amir (NMW 61472, 46081); Scaphiodon niger (NMW 51655, 51654, 51656); Scaphiodon saadii (NMW 51666).

Iranian material: CMNFI 1993-0154, 1, ? mm standard length, Markazi, Sharra River near Far (34º03'N, 49º20'E); CMNFI 1995-0020, , mm standard length, (); CMNFI 1995-0021, , mm standard length, (); CMNFI 1997-0004, , mm standard length, ();

Comparative material: BM(NH) 1974.2.22:1856, 227.1 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1934.9.5:3-5, 6, 14.8-45.5 mm standard length, Rawanduz River, Razanok.

Capoeta fusca
Nikol'skii, 1897

Courtesy of S. A. Johari
Courtesy of S. A. Johari

Common names

سياه ماهي (= siah mahi, meaning black fish).

Systematics

The 2 syntypes, listed in Latin as from "Mondechi in Persia orientali", are in the Zoological Institute, St. Petersburg (ZISP 11108) and measure 121.9-172.9 mm standard length. Berg (1949) gives the locality in Russian as "Mondekhi, northern periphery of the Bajistan Salt Desert in southeast Khorasan". This locality is possibly Mandehi or Miandehi at 34°53'N, 58°38'E. Nikol'skii (1897) lists a series of specimens in Latin, presumably all of which he regarded as types, sic:- "11108. Mondechi in Persia orientali. 12.IV.96 (2). 11109. Persia orientalis. 1896. (6). 11110. Persia orientalis. 1896. (5). 11111. Persia orientalis. 1896. 11112. Kuss in Persia orientali. 6.IV.96.", the last two lacking number of specimens. Berg (1949) gives 20 specimens for 11109, 6 specimens for 11110, and 1 specimen for each of the last two. Catalogue dates in ZISP for all these are 26.IV.96, presumably new style, while Berg (1949) gives new style dates 24.IV.1896 for the first and 18.IV.1896 for the last (and this last is 26.IV.1896 in the catalogue). Only ZISP 11108 specimens are regarded as syntypes by Berg (1949). Berg (1949) also points out the confusion over the date when Zarudnyi, the collector, was at "Kuss" (= Khusf at 32°46'N, 58°53'E) given by Nikol'skii as 6.IV.96 old style but on this date Zarudnyi was at "Kiaz-khak" near Asadabad (35°38'N, 59°21'E) south of Mashhad and only reached Khusf on 8 (or 20 new style).VI.96. This is not particularly critical in this instance but serves to point out the difficulties of reconciling literature, field notes, catalogues, and jar labels.

Capoeta nudiventris Nikol'skii, 1897 is a synonym. The syntypes are in the Zoological Institute, St. Petersburg (ZISP 11106) according to Berg (1949) and comprise 3 fish 92.4-121.5 mm standard length. Berg (1949) gives the type locality as "Zeride near Bajistan in southeast Khorasan, 30.IV.1896" (the date in the jar is 26.IV.1896). Nikol'skii (1897) lists 3 collections all from "Saride in Persia orientali. 18.IV.96." with numbers 11105, 11105 (presumably an error for 11106), and 11107 and 6 (actually 7 in the jar and according to Berg (1949)), 3, and 5 specimens respectively. Berg (1949) lists the 5 specimens under 11107 as from "Chakhak in the Al'kor region between Bajistan and Birjand. 9.V.1896", presumably at 33°17'N, 58°54'E. These 5 fish are 37.0-55.2 mm standard length, collected on 25.IV.1896 in the ZISP catalogue and not listed as types in the jar, nor in the catalogue, nor in Berg (1949). The 7 fish in ZISP 11105 measure 46.8-75.3 mm standard length, are from the same locality listed under ZISP 11106 in Berg (1949) and are listed as types in the ZISP catalogue, though not in Berg (1949). Judging from the labels and catalogue sheets, the types are probably from Sarideh at 34°22'N, 58°14'E and comprise 11105 and 11106.

Rainboth (1981) places both fusca and nudiventris in the genus Schizocypris on the basis of the enlarged scales around the vent and anal fin base, a condition reported on by Berg (1949) also but not considered by this latter author to warrant inclusion of these fish in Schizocypris.

Key characters

The strong mode of 7 branched dorsal fin rays, distribution, and the relatively low scale count aid in identifying this species.

Morphology

Dorsal fin with 3 unbranched rays and 7-8, strong mode at 7, branched rays, anal fin with 3 unbranched and 5 branched rays, pectoral fin branched rays 14-20 and pelvic fin branched rays 7-9. Lateral line scales 40-62, mostly 47-56. Scales are found regularly arranged over the whole body and are enlarged around the anus and anal fin base. There is a pelvic axillary scale. Scales are oval and have a subcentral, markedly anterior focus, numerous radii on all fields and moderate numbers of circuli. Gill rakers 11-20, short and touching the raker below when appressed. The mouth is horseshoe-shaped. The last unbranched dorsal fin ray is weak with only a few fine denticles along the basal half. The pharyngeal teeth are very spatulate up to the tip but are thick. There is an occasional trace of a fifth tooth in the major row but all the fish examined had only 4 strongly developed main row teeth. The gut is very elongate with several anterior and posterior loops.

Some populations or individuals may show a very light belly extending up onto the lower flank rendering scales hard to see. Capoeta nudiventris was apparently founded on specimens like this. Some scales low on the flank are incompletely imbricate and deeply embedded in the skin. Berg (1949) in examining the types of fusca and nudiventris found the extent of the scales ventrally to be the same and nudiventris is not naked on the lower flank and belly.

Meristics for Iranian specimens:- dorsal fin branched rays 7(77); anal fin branched rays 5(77); pectoral fin branched rays 14(1), 15(1), 16(8), 17(23), 18(26), 19(13) or 20(5); pelvic fin branched rays 7(8), 8(64) or 9(5); lateral line scales 46(4), 47(6), 48(8), 49(10), 50(10), 51(9), 52(9), 53(9), 54(9), 55(2) or 56(1); total gill rakers 13(1), 14(11), 15(25), 16(26), 17(11), 18(1), or 20(1); pharyngeal teeth 2,3,4-4,3,2(20); and total vertebrae 40(9), 41(42), 42(20) or 43(4).

Sexual dimorphism

Unknown but males presumably bear large tubercles in the breeding season.

Colour

The back and flanks are dark while below the lateral line the body can be very light. The dorsal, anal and caudal fin membranes are dark. Young fish may have a mid-lateral stripe as wide as the eye ending in an indistinct dark blotch on the caudal fin base. The peritoneum is dark brown to black.

Size

Reaches 21.5 cm total length (Johari et al., 2009).

Distribution

This species is found in eastern Iran in the Tedzhen River (including Kashaf River), Dasht-e Kavir, Bejestan, Dasht-e Lut and Sistan basins in rivers, springs and qanats, some of the latter not easily located on maps (Nikol'skii, 1899; Berg, 1949; Abdoli, 2000). A record from the "Schalman Rud" presumably in the Caspian Sea basin is most probably an error (Wossughi, 1978). Johari et al. (2009, 2010) record this species from the Ghoorghoori, Asafshad, Mardan Shah, Gazdmoo and Afin rivers in Qae'nat province and in 44 qanats of Birjand County in eastern Iran.

--- Zirkhuch may be Zir-e Kuh at 32°48'N, 59°50'E ? check this and if reasonable search for Zir and add in all text - Zirkhuch is in eastern or southeastern Khorasan for sure

Zoogeography

Saadati (1977) considers that this species entered eastern Iran from the west via the Namak Lake basin. See also above under genus.

Habitat

Karaman (1969) considers that this species shows the greatest adaptation among Capoeta species to desert life: an elongate and low body, scaleless belly in many individuals, weak spiny dorsal fin ray, reduced number of dorsal fin rays, short dorsal fin which can easily lie flat against the body, and the mouth structure. Johari et al. (2009) studied 10 qanats in Birjand County and found the following ranges: 3.8-24.9ºC, 0-6.3 p.p.t. salinity, 7.7-8.5 pH, 3.8-1164 μS, 6.3-13.8 mg/l dissolved oxygen, 0.31-11.5 mg/l nitrate, 0-0.8 mg/l nitrite, 0.04-0.29 mg/l ammonia, 185-750 mg/l total hardness, 2.17-815 total dissolved solids, 25-410 mg/l calcium, 0-100 mg/l magnesium, 0.16-340 mg/l sulphate, 2.3-27 mg/l potassium, 0.01-0.14 mg/l chlorine and 0.2-0.95 mg/l phosphate. No mortalities were noted in fish kept in salinities up to 10 p.p.t. for 120 hours, but higher levels started to show progressive mortalities. As salinity increased, fish became darker and dead fish were almost black. The fish exhibited schooling behaviour both in aquaria and in their natural environment.

Age and growth

Johari et al. (2009) found a total length/weight relationship of body weight = 0.01010 x TL2.9477 for 600 fish  from 10 qanats in Birjand County. Patimar and Mohammadzadeh (2011) examined fish from the Shadmehr qanat in south Khorasan and found a maximum age of 5+ years, negative allometric growth for males and isometric for females, males grew faster than females, and von Bertalanffy growth models Lt=18.74(1-e-0.33(t+0.473)) for males and Lt=22.35(1-e-0.32(t+0.333)) for females.

Food

Gut contents of the few fish examined contained fragments of large plants including large seeds, filamentous algae and sand grains. Johari et al. (2009) found this species to be herbivorous based on relative gut length and to be relatively gluttonous based on gut vacuity index. Large plants and filamentous algae made up 86.8% of the food but molluscs, aquatic insects and frog eggs were secondary foods. Feeding was highest in December and January before spawning and in August and September when presumably productivity was greatest. In March to May, the spawning season, feeding was reduced. Badri Fariman et al. (2011) examined fish from the Birjand area and found a food preference of 86.6 for plant material and 13 for benthos.

Reproduction

Fish caught in April and May have mature eggs along with some immature eggs, indicating that spawning may occur in stages. Fish caught in November have small but obvious and developing eggs. Johari et al. (2009) found the reproduction period began in March and lasted until the latter part of May based on the gonadosomatic index. Patimar and Mohammadzadeh (2011) found a sex ratio of 1:2.42 in favour of females for their south Khorasan fish, with reproduction in the qanat between May and August with the gonadosomatic index highest for males in June and for females in July. egg diameters attained 2.05 mm, maximum fecundity attained 22,773 eggs and relative fecundity up to 583 eggs/g. Badri Fariman et al. (2011) also found March to be the spawning time based on the gonadosomatic index, but with fry found at different lengths during all sampling times, spawning may occur more than once a year.

Parasites and predators

Black spots on the head and fins (syntypes of nudiventris as noted by Nikol'skii (1897)) were probably encysted larvae of trematodes (Berg, 1949). Johari et al. (2009) found the trematode Clinostomum in various body parts and their qanat fishes showed lordosis and scoliosis.

Economic importance

This species has been studied in aquaria for the toxicity of lead acetate (Omidi et al., 2009). Toxicity decreased with increase in water hardness, qanat water with a high water hardness (310 mg L-1) showing low toxicity. Mansouri et al. (2011) found that cobalt accumulates in tissues of this species under experimental conditions, with liver showing the most and gills the fastest elimination. Mansouri et al. (2011) examined fish from qanats in the Birjand region for heavy metals (nickel, cobalt, cadmium, chromium and zinc) finding the highest levels in liver and lowest in gills and variations in levels between sample sites. All levels were acceptable in human health consumption terms.

It will feed on mosquito larvae under aquarium conditions and could have been a better candidate for combating malarial mosquitos than the exotic and deleterious Gambusia holbrooki.

Conservation

A widely distributed species apparently able to survive in a wide range of desert habitats, it may not be in need of conservation.

Further work

Biology in mostly unknown and would help confirm the impression that it is not in need of conservation.

Sources

Type material: See above, Capoeta fusca (ZISP 11108) and Capoeta nudiventris (ZISP 11105, 11106).

Iranian material: CMNFI 2007-0005, 7, 27.8-84.2 mm standard length, Semnan, spring at Nardin (ca. 37º03'N, ca. 55º47'E); check ID? CMNFI 2007-0015, 8, 60.1-85.6 mm standard length, Khorasan, qanat at Khalaj (ca. 34º54'N, ca. 58º52'E; CMNFI 2007-0016, 8, 85.5-171.4 mm standard length, Khorasan, qanat and jube at Bidokht (ca. 34º21'N, ca. 58º46'E); CMNFI 2007-0017, ?, ? mm standard length, Khorasan, qanat at Dasht-e Bayaz (ca. 34º02'N, ca. 58º47'E); CMNFI 2007-0018, 15, 21.7-92.4 mm standard length, Khorasan, Shur River (ca. 33º52'N, ca. 59º41'E); CMNFI 2007-0019, 9, 32.7-141.3 mm standard length, Khorasan, qanat between Esfideh and Abbasabad (ca. 33º29-39'N, ca. 59º38-46'E); CMNFI 2007-0020, 23, 43.7-115.1 mm standard length, Khorasan, qanats at Marak and Rabi'an (ca. 32º55-58'N, ca. 59º26-27'E); CMNFI 2007-0021, 16, 24.8-56.3 mm standard length, Khorasan, Shah Abbas qanat in Asadabad (32º55'N, 60º01'E); CMNFI 2007-0022, 6, 56.7-112.1 mm standard length, Khorasan, qanat pool at Mud-e Dahanab (32º43'N, 59º31'E); CMNFI 2007-0023, 6, 82.5-113.1 mm standard length, Khorasan, qanat at Sarbisheh (32º34'N, 59º48'E); CMNFI 2008-0201, 1, ? mm standard length, Khorasan, Birjand qanats (32º52'N, 59º12'E); CMNFI 2002-0202, 1, ? mm standard length, Khorasan, Birjand qanats (32º52'N, 59º12'E); BM(NH) 1958.11.7:1-6, 6, 26.1-90.9 mm standard length, Khorasan, near Jajarm (no other locality data).

Capoeta trutta
(Heckel, 1843)

Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi



Courtesy of H. R. Esmaeili

Common names

tu'ini (and variant spellings in transliteration such as touyeni, tuyeni, tuini or too'ini) in Khuzestan (meaning unknown); tu'ini gelkhorak in Khuzestan (see C. damascina for meaning); shir mahi (= milk fish), barg bidy or barg-e bidi (= willow leaf, perhaps from shape and colour), berzem.

[twena, hemira, tela morqat, tela moraqqat; ethra at Mosul (Heckel (1843b), or takal handscherli (takal = soft or flexible presumably from its small scales, handscherli = armed with a dagger or knife from the dorsal fin spine) at Aleppo (Heckel, 1843b), all in Arabic; trout barb].

Systematics

Rainboth (1981) places this species in Schizocypris on the basis of enlarged scales forming a split to encompass the urogenital region and a bare to partially bare mid-dorsal strip anterior to the dorsal fin. However the schizothoracine fishes are quite different (see accounts for Schizothorax, Schizopygopsis and Schizocypris) and this placement is not accepted here.

The type localities of Capoeta Trutta as given by Heckel (1843b) are "Gewässern bei Aleppo" and the "Tigris bei Mossul". The syntypes are in the Naturhistorisches Museum Wien according to Krupp (1985c) as follows: NMW 55935-37, 55942, 6 specimens 94-274 mm standard length from Mosul, NMW 55926, 55928, 55940-41, 7, 68-192 mm standard length from Aleppo, and in the Senckenberg Museum Frankfurt (SMF 2567 (formerly NMW), 1, 407 mm standard length, from Mosul and SMF 923 (formerly NMW), 1, 175 mm standard length, from Aleppo. Four other syntypes are under NMW 55939, 1 other syntype under NMW 55938 and a dried syntype under NMW 58875. Eschmeyer et al. (1996) lists similar material with the numbers of fish under each catalogue number detailed thus: NMW 55926 (1), NMW 55928 (2), NMW 55935-37 (2, 2, 1), NMW 55939-42 (4, 1, 3, 1), possibly 1 fish in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 3164, formerly NMW), 1 syntype in the Senckenberg Museum Frankfurt (SMF 923, formerly NMW) and 1 syntype SMF 2567 (formerly NMW), and 1 dried syntype from the Museum für Naturkunde, Universität Humboldt, Berlin (ZMB 8789; not located in February 2006). The catalogue in Vienna lists only 5 specimens although the card index in 1997 lists NMW fish as syntypes in agreement with Eschmeyer et al. (1996).

Key characters

The combination of small scales, transverse mouth, dorsal and anal fin branched ray counts, the very strong last unbranched dorsal fin ray (longer than head length - usually strong but rarely weak), and the colour pattern identifies this species.

Morphology

Dorsal fin with 3-5 unbranched rays followed by 7-9, usually 8, branched rays, anal fin with 2-3 unbranched rays followed by 5 branched rays, pectoral fin branched rays 14-18, and branched pelvic fin rays 7-8. Hanel et al. (1992) found 23-31 denticles or teeth on the serrated dorsal fin ray, the largest near the centre of the ray length. Scales in lateral line 68-90, scales above lateral line 15-18 and scales below lateral line 10-17. The back anterior to the dorsal fin is compressed and lacks scales except near the occiput. Scales have a protruding anterior margin but are otherwise rounded, anterior and posterior radii, fine circuli and a subcentral anterior focus. There is a pelvic axillary scale. Gill rakers 23-33, on the lower arm 18-25 (with lowest counts in smallest fish). The rakers reach the second raker below when appressed. Pharyngeal teeth 2,3,4-4,3,2. Teeth are broadly spoon-shaped at the tip, with narrow cusps and stems such that they are quite fragile. A frequency distribution of counts was not taken because of this fragility. Total vertebrae 43-46. The mouth is inferior and transverse with a strong horny cover to the lower jaw. The gut is very elongate with numerous anterior and posterior loops. The karyotype of fish from the Tigris River of Turkey is 2n=150, possibly hexaploid, with 35 meta-submetacentric chromosomes, 40 pairs of subtelo-acrocentric chromosomes with NF=220 (Kılıç Demirok and Ünlü, 2001).

Meristics for Iranian specimens:- branched dorsal fin rays 8(34); branched anal fin rays 5(34); branched pectoral fin rays 14(1), 15(8), 16(18), 17(6) or 18(1); branched pelvic fin rays 7(32) or 8(3); lateral line scales 68(2), 69(1), 70(1), 71(4), 72(5), 73(3), 74(5), 75(2), 76(3), 78(3), 79(1), 80(1), 81(1), 83(1) or 84(1); total gill rakers 22(1), 24(5), 25(4), 26(3), 27(7), 28(8), 29(3), 30(2) or 31(1); and total vertebrae ?more 43(1), 44(6), 45(3) or 46(2) - NMC 79-269, 367, 384, 269, 268, Behnke 231 done.

Sexual dimorphism

Males bear a single tubercle on each flank scale, sometimes 2 tubercles, positioned about the middle of the exposed scale or nearer the posterior edge. The head has small and widely scattered tubercles on the top and sides and large tubercles around the snout from eye to eye below the nostril level. Large tubercles occur in single files on the anal and dorsal fin rays, particularly the posterior rays, becoming apparent on the more anterior rays as tuberculation develops more highly.

Colour

The head and body and the dorsal fin (and sometimes the caudal fin) are covered with small, distinctive black spots, often c- or x-shaped. Spots are apparent through the silver flank colour. Some fish in Khuzestan lack spots but transitional specimens from fully spotted through weakly spotted to immaculate are found. Colour is brownish to yellowish or olive-green on the back with silvery-white flanks and the belly lighter, white with silvery tints. Some fish are very pale almost whitish. Upper flank scales in particular are outlined with dark pigment. The eye is orange above or mostly silvery. Lower fins are orange to yellow at the base and blackish distally, or may be orange to yellow overall. The dorsal and caudal fins are grey or hyaline. The lower rays of the caudal fin have a slight orange-yellow tint. The peritoneum is dark brown to black.

Size

Attains at least 45.8 cm total length. Heckel (1843b) gives 1 Schuh 8 Zoll, or 52.7 cm.

Distribution

Found in the Quwayq, Orontes and Tigris-Euphrates basins including the Iranian portion of the latter at such places as Dez River near Dezful, Gheshlagh Dam Lake, Sanandaj - others? (Berg, 1949; Marammazi, 1995; K. Abbasi, see photograph above; Bahrami Kamangar et al., 2012; Oğuz et al., 2012) and the Gulf basin in the Zohreh River.

upper Mand ? to check on maps

Zoogeography

Its relationships with other Capoeta species is generally unclear, as are the larger zoogeographical relationships of this and other Capoeta species. See also above under genus.

Habitat

Marammazi (1994) considers this species to be stenohaline but nonetheless more widely distributed than stenohaline Barbus (= Mesopotamichthys) sharpeyi in the Zohreh River which drains to the northern Persian Gulf.

Age and growth

The majority of the population studied by Ünlü (1991) in the Tigris River in Turkey are in age groups 2 and 3 although males live to age 7 and females age 10. Females are usually longer and heavier than males of the same age. Males comprise 41.26% and females 58.74% of this population. In a stream in the Euphrates River drainage of Turkey, Gul et al. (1996) found fish to live for 8 years with 60-90% of the fish in age groups 1 to 3. Females comprised 53.3% and males 46.7% of the population. Kalkan (2008) studied a population in the Karakaya Dam lake on the Turkish Euphrates River. Maximum age was 7 years, age groups 4 and 6 were mostly females whereas age group 3 was mostly male, age-length, age-weight and length-weight formulae were given, and the average growth condition factor was 1.30 for females and 1.28 for males.

Patimar and Farzi (2011) observed a maximum age of 6+ years in the Meymeh River of western Iran. The sex ratio was 1:1.35 for male:female and length-weight relationships were TL = 0.0266TW2.7134 for males, TL = 0.0258TW2.7251 for females and TL = 0.026TW2.7217  for the population. von Bertalanffy parameters were L = 45.86 mm, k = 0.14 year-1, t0 = -1.15 and =5.68 for males, 50.79, 0.13, -1.45 and 5.81 for females, and 48.6, 013, -1.28 and 5.72 for the population.

Food

Gut contents include diatoms, green algae and large amounts of sand.

Reproduction

Spawning in both the Tigris and Euphrates rivers in Turkey took place in May-June. Males mature at age 2 and females at age 3 in both rivers. Ripe egg size in the Tigris varied between 1.33 and 2.11 mm and egg numbers between 4713 and 18240. Ripe eggs in the Euphrates attained 1.04 mm and the maximum number of eggs per gramme of gonads was 666. Fish from Khuzestan had well-developed eggs on 30 January while adult fish taken on 7 July were not in reproductive condition.

In Iran fish spawn in March-May (Abdoli, 2000). Fish from the Meymeh River examined by Patimar and Farzi (2011) spawned in these months and had a maximum egg diameter of 1.9 mm, a mean absolute fecundity of 7594 eggs and a mean relative fecundity of 70 eggs g-1 body weight.

Parasites and predators

Molnár and Jalali (1992) report the monogenean Dactylogyrus pulcher from this species in the Dez River of Khuzestan. Gussev et al. (1993a) describe a new species of monogenean from this species in the Dez River, Dactylogyrus microcirrus. Baska and Masoumian (1996) describe two new species of Myxosporea from fish caught in the Karun River at Ahvaz, Myxobolus molnari taken from the gills and Myxobolus mokhayeri taken from between the soft rays of the fins. The latter species is named after Dr. Baba Mokhayer, an internationally renowned Iranian professor. The new species are of minor pathological importance as the infections are of low intensity and prevalence. Masoumian and Pazooki (1999) list Myxobolus molnari and M. mokhayeri from this species from localities in Khuzestan. Peyghan et al. (2001) record Neoechinorhynchus sp. and Rhabdocona sp. from fish from Khorramabad rivers. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. and Tracheliastes polycolpus on this species. Oğuz et al. (2012) record the acanthocephalan Neoechinorhynchus zabensis from fish in the Dez River.

Economic importance  

Duman and Duman (1996) give the nutritional value of Capoeta trutta from Keban Dam Lake in Turkey but this fish is little used in Iran. However, Peyghan et al. (2001) report that is is an economically important species with a good market value in the Khorramabad region.

Conservation

This species does not appear in need of conservation but its biology is too poorly known in Iran to be certain. Bahrami Kanagar et al. (2012) provided baseline haematological and biochemical indices for this species in Gheshlagh Dam Lake, Sanandaj that could be used in health monitoring. Kalkan (2008) recommended prohibition of fishing in Turkey during March-August and fish under 22.62 cm should not be retained.

Further work

The biology of this species and its relationships to other Capoeta species needs work.

Sources

Type material: ?

Iranian material:- CMNFI 1979-0020, ?, ? mm standard length, (); CMNFI 1979-0268, 3, 115.7-141.2 mm standard length, Lorestan, between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0269, 2, 114.1-144.1 mm standard length, Lorestan, between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0367, 2, 29.7-54.1 mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E); CMNFI 1979-0368, 8, 36.3-67.9 mm standard length Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0376, 1, 55.2 mm standard length, Khuzestan, river tributary to Karkheh River (32º48'30"N, 48º04'30"E); CMNFI 1979-0384, 1, 218.4 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1991-0153, 2, 153.8-217.2 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1995-0020, ?, ? mm standard length, (); CMNFI 1995-0021, ?, ? mm standard length, (); CMNFI 1995-0030, ?, ? mm standard length, (); CMNFI 2007-0100, 1, 136.7 mm standard length, Azarbayjan-e Gharbi, Kalwi Chay near Piranshahr (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0109, 11, 61.3-167.4 mm standard length, Kordestan, Qeshlaq River basin north of Sanandaj (ca. 35º33'N, ca. 47º08'E); CMNFI 2007-0110, 3, 96.6-160.3 mm standard length, Kordestan, Yuzidar River basin (ca. 35º05'N, ca. 46º56'E); CMNFI 2007-0113, 1, 74.8 mm standard length, Kermanshahan, Razavar River 35 km northwest of Kermanahah (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0116, 1, 95.9 mm standard length, Kermanshahan, Gav Masiab west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, 2, 153.8-217.2 mm standard length, Kermanshahan, Gav Masiab near Sahneh (ca. 34º24'N, ca. 47º40'E); ZMH 2511, 1, 319.0 mm standard length, Kermanshahan, Karasu-Gamasiab-Seymarreh (no other locality data).

Comparative material: BM(NH) 1931.12.21:8, 1, 113.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1968.12.13:376-390, 15, 35.6-123.3 mm standard length, Syria, Euphrates River at Mayadine (?); BM(NH) 1974.2.22:1374-1377, 4, 66.3-91.2 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1382, 1, 86.1 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E); BM(NH) 1974.2.22:1388-1389, 2, 259.4-273.3 mm standard length, Iraq, Tigris River at Samarra (?).

Genus Capoetobrama
Berg, 1916

Capoetobrama kuschakewitschi
(Kessler, 1872)

This species is reported from the Karakum Canal of Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually be found in the Tedzhen River and Caspian Sea basins of Iran. No Iranian record.

Genus Carasobarbus
Karaman, 1971

?

Some of the past literature on this genus appeared under Barbus (q.v.)

 

Carasobarbus luteus
(Heckel, 1843)



Kol River, courtesy of H. R. Esmaeili

Common names

حمري (= hemri), himri; sangal or zangol (= blackish, used at Kermanshah, J. Valiallahi, pers. comm., 2001); lab matiki (= from lipstick by professional fishermen at Kermanshah in reference to red lips, from J. Valiallahi, pers. comm., 2001).

[himri, hamria, hamra, binni hamour, binni hamri, bunni himri, binni, binni shifatha, beni asphar (= yellow son), beni abjad (= white son), beni hamra (= red or yellow son), zuri or bartema, all in Arabic; golden barb, yellow barbel].

Systematics

Heckel (1843b) gives localities for the types of Systomus luteus as "Orontes", and "Tigris", and in the next sentence at "Aleppo" and "Mossul". Two syntypes were examined in the Naturhistorisches Museum Wien under NMW 54250 (but see below). Krupp (1985c) records a 301 mm standard length syntype from Aleppo formerly in the Naturhistorisches Museum Wien, now in the Senckenberg Museum Frankfurt as SMF 6784. Eschmeyer et al. (1996) list the following syntypes: NMW 10827 (1 fish), NMW 54247 (2), NMW 54248 (1), NMW 54249 (1), NMW 54253 (2), NMW 54254 (3), NMW 54255 (2), NMW 54520 (2), NMW 80043 (2) and possibly 2 syntypes in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2463, formerly NMW) as well as the syntype in Frankfurt. The catalogue in Vienna seems to list 5 specimens but this part of the catalogue is overwritten and difficult to interpret. The card index in 1997 lists NMW 53680a (1 fish, the lectotype), 53674 (1), 53675 (1) and 53676 (1) as the syntype series.

Systomus albus Heckel, 1843 from the "Tigris" and "Orontes" and Systomus albus var. alpina Heckel, 1847 are synonyms.

Systomus albus var. alpina was described from the "Flusse Kara-Agatsch und den Alpenseen Pire-San und Deria Kaserun" (= Qarah Aqaj River and Lake Famur, Fars; Pire-San being Parishan and Deria Kaserun being Lake Kazerun, both other names for Lake Famur) (Heckel, 1847b). Krupp (1985c) records 4 syntypes of alpina from Shiraz (sic), Th. Kotschy as NMW 53679 (2 fish) and NMW 53681 (2). NMW 53678 (5 fish, 27.6-60.8 mm standard length), NMW 53679 (2 fish, 63.8-70.5 mm standard length), and NMW 53681 (2 fish, 79.6-93.3 mm standard length) are from the "Kara Agatsch bei Schiraz"; and NMW 53682 (2 fish, 201.7-203.7 mm standard length) are from the "Alpenseen Pire-san und Deria Kaserun": all are possibly syntypes of Systomus albus var. alpina although the catalogue in Vienna lists 5 fish under this name in one column and 4 fish in smaller writing in the adjacent column. The card index in 1997 lists syntypes under NMW 53678 (5), 53679 (2), 53681 (2) and 53682 (2, one of which is the lectotype). Eschmeyer et al. (1996) list 2 fish in the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH 2464) as possible former NMW types of this taxon.

A dried specimen of Systomus albus from Mosul collected by Th. Kotschy may be a syntype (NMW 59485). Eschmeyer et al. (1996) gives the syntypes of this species as NMW 53674 (1), NMW 53675 (1), NMW 53676 (1), NMW 53677 (1), NMW 53680 (1), NMW 91400 (1, dry) and SMF 812 (1), formerly NMW. Krupp (1985c) records the syntype of albus in the Senckenberg Museum Frankfurt under SMF 812 as being 84 mm standard length. The Vienna catalogue lists 4 fish under Systomus albus but the card index in 1997 lists the same NMW fish as Eschmeyer et al. (1996) as above with NMW 53680 as lectotype.

Barbus parieschanica Wossughi, Khoshzahmat and Etemadfar, 1982 is presumably also from Lake Famur or Parishan judging by the name and is a synonym (note that the species name is first spelt parschanica on page 23 in the abstract in Farsi and on page 44 in the English abstract but in the text species description (page 34) and in the table (page 37) it appears as parieschanica, and this is presumably the intended correct spelling). The species locality in the text is "Noorabad of Mamasany". ?

Saadati (1977) refers to a new and undescribed Cyprinion species from Lar in southern Iran but the fish are Carasobarbus luteus.

Günther (1874) placed this species in Barynotus Günther, 1868, a genus with the type species from West Africa. Barynotus is preoccupied in Coleoptera and was replaced by Barbellion Whitley, 1931 (Eschmeyer, 1990). Most authors place the species in Barbus although Karaman (1971) erected a new genus for it, Carasobarbus; and Krupp (1985c) also synonymises Carasobarbus with Barbus. Bănărescu (1997) and Ekmekçi and Banarescu (1998) recognise Carasobarbus as a valid genus however. Borkenhagen et al. (2011) recognise C. luteus as a single, generalist species tolerating a wide variety of habitats.

A group of related species share characters with this species (see also under Kosswigobarbus kosswigi). Carasobarbus may be the generic names for certain members of the group.

Key characters

This species is characterised by a low scale count, smooth last unbranched dorsal fin ray, one or two pairs of barbels, and 10 branched dorsal and 6 branched anal fin rays.

Morphology

Dorsal fin with 4 unbranched rays followed by 9-11, usually and modally 10, branched rays. The last unbranched dorsal fin ray is smooth, thickened, sharp-edged and spine-like. Anal fin with 3 unbranched rays followed by 5-7, usually and modally 6, branched rays. Pectoral fin branched rays 13-17 and pelvic fin branched rays 7-9, usually 8. Lateral line scales 23-36. There is a pelvic axillary scale. There are moderate to many anterior field radii and many posterior field radii and occasionally few lateral radii. The focus is central to subcentral anterior, the anterior scale margin is wavy and the exposed part of the scale is coarse. The concealed part of the scale has numerous fine circuli. Total gill rakers 7-14, reaching the adjacent raker when appressed, sometimes forked at the tip and with spinules on the anterior side. Pharyngeal teeth usually 2,3,5-5,3,2, with the anterior 2-3 teeth rounded and heavier than the posterior teeth. Variants may have 2,3,4 or 1,3,5 (Borkenhagen, 2005). Posterior teeth are hooked at the tip and the grinding surface below the tip is irregular with a protuberant knob which may be striated. The gut is elongate with both posterior and anterior loops.

The mouth is terminal to subterminal and lips are weakly developed. There is one pair of short and thin barbels at the corner in most descriptions. Number and frequencies for 130 fish are 2 barbels (47 or 36.2%), 3 barbels with left anterior present (7 or 5.4%), 3 barbels with right anterior present (5 or 3.8%), or 4 barbels (71 or 54.6%). However, this sample is 112 fish or 86.2% from Fars and Hormozgan. Fish from these provinces, at such localities as the lower Mand River and the Sar Khun oasis north of Bandar Abbas consistently have a high frequency of 4 barbels (58.9%), and with 3 barbel counts included 68.8%, than fish from the Tigris River basin. Even the 18 fish from the Tigris River basin had 5 fish with 4 barbels so, at least in the eastern part of this species range, 4-barbelled fish are not rare.

Body form varies with habitat (Ali, 1982a), there being lake and river forms as with many other cyprinid species.

Iranian specimens have the following meristics: dorsal fin branched rays 9(7), 10(102) or 11(7); anal fin branched rays 5(3) or 6(114); pectoral fin branched rays 14(12), 15(44), 16(48) or 17(13); pelvic fin branched rays 7(9), 8(107) or 9(1); lateral line scales 23(2), 24(10), 25(20), 26(22), 27(28), 28(16), 29(14), 30(4) or 31(1); total gill rakers 8(6), 9(24), 10(40), 11(28), 12(12), 13(3) or 14(2); pharyngeal teeth 2,3,5-5,3,2(19), 2,3,4-5,3,2(4) or 2,3,5-4,3,2(2); and total vertebrae 36(8), 37(53), 38(70), 39(25) or 40(1).

Sexual dimorphism

A 12.7 cm specimen from the Mand River has tubercles on the dorsal, anal, caudal, pectoral and pelvic fins, most strongly on the anal fin rays. Fine tubercles cover the top and sides of the head. A 20 cm fish from the same collection lacked tubercles. Another fish from the lower Mand River (128.5 mm standard length) also has fine tubercles on the upper flank scales as well as the head and fin rays. Ali (1982) reports no sexual dimorphism for Iraqi fish.

Colour

The back and upper flank is dark brown, greenish black or grey-green fading to a whitish or silvery belly all overlain by an orange to yellowish tinge. On the upper flank, scale bases are black-brown with a light blue-grey margin. There is a dark stripe along the mid-line of the back and a dark mid-lateral stripe. Fins are greyish to lime-green, reddish-yellow or orange, becoming blackish distally. The pectoral and pelvic fins tend to be more orange than the anal and caudal fins which are more a faint lime-green. The lips are orange. The eye rim is yellow-green. The peritoneum is black. Small fish have a collection of melanophores at the mid-base of the caudal fin forming a spot-like structure.

The fish described by Heckel (1847b) as Systomus albus var. alpina were also painted live and had a lead-grey body, light brown at the head and reddish-white on the belly. Each scale was black-brown at the base and light blue-grey at the margin, particularly on the upper flank. All fins were blackish and the eyes orange-red.

Size

Attains 38 cm calculated maximum length and 501 g (Ahmed, 1982) or 750 g (Borkenhagen, 2005). Heckel (1843b) gives 17 Zoll for Systomus albus (= 44.8 cm).

Distribution

This species is found in the Orontes and Quwayq rivers and the Tigris-Euphrates basin. In Iran, it is found in the Tigris River basin including the Hawr Al Azim marsh, the Gulf basin including the Helleh, Dalaki, Shapur, Mand and Dasht-e Palang rivers and Lake Famur, the Lake Maharlu basin, the Hormuz basin and the Kor River basin (Wossughi, 1978; Bianco and Banarescu, 1982; Gh. Izadpanahi, pers. comm., 1995; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000). The record from the Kor River basin (Abdoli, 2000) needs confirmation with specimens.

Zoogeography

Karaman (1971) considers that the closest relatives of this species were to be found in India and southern Asia.

Habitat

van den Eelaart (1954) reports that this species in Iraq is a resident in still water and the slower sections of rivers and is the main fish in canals. In summer it goes to the deeper basins of marshes and remains in the shade of plants. It tolerates warm water but does not go into open waters. Al-Hassan and Muhsin (1986) record this species from the Khor al Zubair in southern Iraq where annual temperature range is 12-30°C and annual salinity change is 28-47‰. The fish appear unaffected by these conditions while Heteropneustes fossilis is moribund. Mohamed et al. (1993) report Barbus (= Carasobarbus) luteus from 2 km southward of Fao, Iraq in a pure marine habitat (temperature 13-35°C and salinity 30-47‰). The fish were caught in April which is the flood season.

Age and growth

Ahmed et al. (1984) studied the reproductive cycle of this species in the Hawr al Hammar in southern Iraq near Basrah. Maturity is attained at a minimum of 11.2 cm for females and 12.2 cm for males, at age 1+. The largest fish are 26.0 cm and age 6. Barak and Mohamed (1983) also found 6 age groups for fish from the Garma Marshes, Iraq. Ahmed (1982) studied a population in Tharthar Reservoir about 65 km northwest of Baghdad and found 7 age groups. This study has the fastest growth of Iraqi populations. Khalaf et al. (1988) worked on a population in a flooded gravel pit about 50 km north of Baghdad in Iraq and found fish up to age group 7+. Growth is greatest in the first year (67 mm) and averaged only 22.5 mm in the following years. Growth is slow in consequence of high salinity (3-6% (sic)) and poor food resources. Mohamed et al. (1993) report fish up to 7 years of age in a marine setting in Iraq, Epler et al. (1996) up to 5+ years in fresh and salty Iraqi lakes. Biro et al. (1988) found fish up to age group 8+ in the Diyala River, Iraq. Al Hazzaa and Hussein (2007) describe larval development and growth in the laboratory using fish from a Syrian hatchery. Gökçek and Akyurt (2008) found fish up to 9 years of age in the Turkish Orontes River and give growth parameters for this population. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 34 Iranian fish measuring 3.20-16.80 cm standard length. The a-value was 0.0232 and the b-value 3.036 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Naama and Muhsen (1986) examined feeding periodicities in this species in the Hawr al Hammar, Iraq. Food is mainly detritus, aquatic plants and algae taken throughout the night and day. Barak and Mohamed (1982) studied food habits in the Garma Marshes, near Basrah, Iraq and found this fish to contain principally aquatic plants, the broken and fragmented leaves and stems of Vallisneria in particular. Diatoms and other algae as well as shrimps, chironomid larvae, gastropods and cladocerans are important foods. Invertebrates are about eight times more important in fish smaller than 30 cm than in larger fish. Plant parts are more important, almost twice as much, in larger fish than smaller. Mohamed et al. (1993) report plant remains to be dominant and fish eggs in lesser quantities in a marine setting in Iraq. Epler et al. (1996) found plants to dominate in fish from fresh and salty Iraqi lakes, although not to the same extent as in Barbus (= Mesopotamichthys) sharpeyi where 95.7-100% of the diet was plants. Tendipedids, worms, detritus and fish were also found in B. luteus.

Khoshzahmat et al. (1981) found that this species did not eat molluscs in Lake Perishan (= Famur), near Kazerun in Iran and assume its diet is aquatic plants.

Reproduction

Spawning in the Hawr al Hammar starts in April and after July no fish are found in a partially spent phase. Eggs are yellow to orange in colour and testes white. The eggs attain 1.86 mm in diameter and number up to 38,433 for the oldest fish. Bhatti and Al-Daham (1978) and Al-Daham and Bhatti (1979) report a spawning season of May-July (peak June-July) for a lower Euphrates River, Iraq population, perhaps as a result of cooler temperatures outside the shallow marshes where warmer temperatures cause an earlier development of gonads. Epler et al. (1996) report spawning in June/July in freshwater Iraqi lakes, earlier in a saline lake. Iranian fish have well-developed eggs in May.

Parasites and predators

Bykhovski (1949) reports a new species of monogenetic trematode, Dactylogyrus persis, from this species in the Karkheh River, Iran. Ebrahimzadeh and Nabawi (1975) list species in the nematode genus Philometra, the protozoan genera Myxosoma and Trypanosoma, the trematode genera Dactylogyrus and Gyrodactylus and the nematode species Camallanus lacustris as well as various unidentified cestodes, trematodes, acanthocephalans and hookworms, from this species in the Karun River. Jalali and Molnár (1990a) records two monogenean species, Dactylogyrus spp., from this species in the Dez River. Molnár and Jalali (1992) describe a new species of monogenean, Dogielius persicus, from this species in the Dez and Karun rivers of Khuzestan. Gussev et al. (1993b) describe a new species, Dactylogyrus carassobarbi, from this species in the Dez River, Khuzestan, the specific name being founded on a misspelling of the genus name Carasobarbus. Masoumian et al. (1994) describe a new species of Myxosporea from the gills of this species in the Karun River, Khuzestan, namely Myxobolus persicus, and later (Masoumian et al., 1996) another new species of Myxosporea, Myxobolus nodulointestinalis, in the gut lining of this species and also from rivers of southwestern Iran. Molnár et al. (1996) report additional new species from this fish in Khuzestan, namely Myxobolus iranicus in the spleen and Myxobolus mesopotamiae in connective tissue of the caudal and pectoral fins. Molnár and Pazooki (1995) record philometrid nematodes from this species in the Karun River, and these are presumed to be a new species.

Masoumian and Pazooki (1999) list Myxobolus persicus, M. karuni, M. sharpeyi, M. nodulointestinalis, M. mesopotamiae and M. iranicus from this species in various localities in Khuzestan. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from Dez River fish. Farahnak et al. (2002) record Anisakis sp. from this fish in Khuzestan Province.

González-Solís et al. (1997) report Proleptinae larvae (Nematoda) from this species in the drainage of Lake Maharlu, Fars. The definitive host is a predatory fish, possibly Mastacembelus mastacembelus, not yet recorded from this basin.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 1.6% with the worm Bothriocephalus opsariichthydis in Khuzestan marshes. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Ergasilus sp., Ergasilus sieboldi and Lernaea sp. on this species.

Economic importance

An important food fish in southern Iraq and Iran (Al-Daham and Bhatti, 1979; Ahmed, 1982). Sharma (1980) reports that hamri were the fourth most important fish species at Basrah fish market, accounting for 267,570 kg from October 1975 to June 1977. Heckel (1847b) reports that they "reach a good size and are very tasty" in Lake Famur, Fars.

In some parts of Southwest Asia this species is regarded as &qut;sacred" kept and bred in special pools where fishing is forbidden (Tortonese, 1934).

The eggs of this species are poisonous (Najafpour and Coad, 2002). A kebab made of about one-quarter of an ovary was eaten. Toxic effects were dizziness, abdominal pain, vomiting, diarrhoea, bitter taste, dryness of mouth, intense thirst, and faintness. One victim as hospitalised for two days and his stomach pumped while a second victim recovered after one day's rest.

Conservation

Vulnerable in Turkey (Fricke et al., 2007). It is a common species in Iranian freshwaters but no detailed conservation assessment has been made.

Further work

The biology of this species in Iran needs study along with its conservation status. Specimens from Fars show differences in body form from those in Khuzestan and this could be investigated.

Sources

Type material: ?

Iranian material: CMNFI 1979-0023, 17, 58.3-161.4 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0024, 1, 61.5 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0026, 2, ? mm standard length, Fars, Shapur River (29º47'N, 51º35'E); CMNFI 1979-0047, 1, ? mm standard length, Fars, Ab-e Paravan (ca. 29º34'N, ca. 52º42'E); CMNFI 1979-0076, 1, ? mm standard length, Fars, Barm-e Shur (29º28'N, 52º41'30"E); CMNFI 1979-0087, 1, ? mm standard length, Khuzestan, Karun River at Ahvaz (31º19'N, 48º42'E); CMNFI 1979-0125, 1, ? mm standard length, Bushehr, Dalaki River near Dalaki (ca. 29º28'N, ca. 51º21'E); CMNFI 1979-0129, 26, ? mm standard length, Fars, spring about 2 km from Farrashband (28º54'N, 52º04'E); CMNFI 1979-0135, 19, ? mm standard length, Fars, Mand River tributary (28º08'N, 53º10'E); CMNFI 1979-0154B, 3, 160.7-258.6 mm standard length, Fars, stream channels at Koorsiah (28º45'30"N, 54º24'E); CMNFI 1979-0155, 2, ? mm standard length, Fars, spring at Gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 6, ? mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0157, 1, ? mm standard length, Fars, qanat at Hadiabad (28º52'N, 54º13'E); CMNFI 1979-0160, 2, ? mm standard length, Fars, spring at Arteshkkadeh Pomp (29º09'N, 53º37'E); CMNFI 1979-0163, 1, 84.9 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0164, 6, 56.6-91.1 mm standard length, Fars, neighbourhood of Shiraz (no other locality data); CMNFI 1979-0187, 31, ? mm standard length, Hormozgan, stream and pools at Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0206, 3, 24.4-25.1 mm standard length, Fars, qanat near Runiz-e Pa'in (29º12'N, 53º40'E); CMNFI 1979-0240, 3, ? mm standard length, Fars, Parishan Lake (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0304, 5, ? mm standard length, Fars, Parishan Lake (ca. 29º31'N, ca. 51º50'E); CMNFI 1979-0347, 2, ? mm standard length, Fars, Pol-e Berengie (29º27'30"N, 52º32'E); CMNFI 1979-0352, 7, ? mm standard length, Khuzestan, marsh in Jarrahi River drainage (30º33'30"N, 48º48'E); CMNFI 1979-0358, 1, 23.7 mm standard length, Khuzestan, pond southeast of Bostan (31º37'N, 48º07'E); CMNFI 1979-0360, 8, ? mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0364, 6, ? mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N, 48º20'30"E); CMNFI 1979-0371, 7, ? mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0687, 7, 124.8-154.1 mm standard length, Fars, Shiraz bazar (no other locality data); CMNFI 1979-0789, 4, ? mm standard length, Fars, Lake Parishan (29º31'N, 51º48'E); CMNFI 1991-0154, 1, ? mm standard length, Khuzestan, Hawr al Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0126, 1, ? mm standard length, Kermanshahan, Sarab-e Yavari (34º28'N, 46º56'E); CMNFI 1993-0127, 1, ? mm standard length, Kermanshahan, Sarab-e Maran (34º44'N, 46º51'E); CMNFI 2007-0060, 2, ? mm standard length, Fars, Chashmeh Ab-e Shirin near Lar (ca. 27º41'N, ca. 54º17'E); CMNFI 2007-0111, 1, ? mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); ZSM 21861, 5, 172.0-217.2 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E).

Comparative material: CMNFI 1987-0017, 3, 97.3-143.9 mm standard length, (); BM(NH) 1934.9.5:6, 1, 117.3 mm standard length, Kurdistan, Ain al Hamra, Shithatha (); BM(NH) 1973.6.21:194, 1, 203.4 mm standard length, Iraq, Shatt al Arab (); BM(NH) 1974.2.22:1338, 1, 134.9 mm standard length, Iraq, Najab Bazar (); BM(NH) 1974.2.22:1346, 1, 108.7 mm standard length, Iraq, Tigris River near Faish Khabour (); BM(NH) 1986.2.14:4-7, 4, 98.6-146.6 mm standard length, Iraq, Baghdad (33º21'N, 44º25'E).

Genus Carassius
Nilsson, 1832

The goldfishes comprise 2-3 species found in Europe, northern Asia and the Far East. Eschmeyer (1990) and Kottelat (1997) comment on the authorship of Carassius. One species is now common in Iran.

These fishes are characterised by a stout and compressed body, last unbranched dorsal and anal fin rays finely serrated, long dorsal and short anal fin, mouth small and terminal, lips thick and fleshy, no barbels, pharyngeal teeth in 1 row and molariform but compressed, numerous gill rakers, and scales large.

Carassius auratus
(Linnaeus, 1758)

Wikimedia Commons
Wikimedia Commons

Carassius auratus from British Fresh-Water Fishes by Rev. William Houghton (1879)
Carassius auratus from British Fresh-Water Fishes by Rev. William Houghton (1879)

Common names

mahi-ye talaee or mahi-ye talai (= gold fish) or اوشين (ooshin or oushein) in Khuzestan; kapur safid by anglers in Khuzestan at Ahvaz; kopur-cheh (= small carp) or كاراس (= karas, karass or karaz) in Mazandaran; kopur cheky (= by the job carp?), kopur chekeh (= drop carp?); ماهي حوض (= mahi-ye howz or mahi-e-hoz, meaning pond or pool fish), mahi-ye howz-e noqrehi (= silvery pond fish, for silvery form), mahi-ye howz-e talaee (= golden pond fish for orange form).

[samak zahabi, buj-buj in Nasiriyah; samti; yayabash in Basrah; karseen in Baghdad; carp thahabi, all in Iraqi Arabic; serebryanyi karas or silver crucian carp in Russian; goldfish for auratus, Prussian carp for gibelio].

Systematics

Cyprinus auratus was originally described from China and Japanese rivers.

Pelz (1987) discusses the scientific name of the goldfish and its confusion with Carassius carassius. All diploid goldfish of western Europe are Carassius auratus auratus (from introductions, presumably including releases and escapes in Iran) and all triploid goldfish are C. auratus gibelio from eastern areas. Bai et al. (2011) also consider triploid and polyploid individuals to be C. auratus gibelio. Goldfish do not appear to be native to Iran but Iranian specimens are sometimes referred to Carassius auratus gibelio (Bloch, 1782) known as the Prussian carp, European goldfish or silver crucian carp. Berg (1948-1949) considers the familiar pet "goldfish" to be a domesticated form of the Prussian carp. However these fish probably have a number of origins - from aquarium stock and from China. Kottelat (1997) tentatively recognises Carassius gibelio (Bloch, 1782) as a species native to eastern Central Europe, and Kottelat and Freyhof (2007) map gibelio as the introduced species in the Caspian Sea basin of Iran. Vasil'eva and Vasil'ev (2000) state that fish named in the literature as Carassius auratus gibelio from Europe, Siberia and eastern Asia are triploids and are not a valid subspecies of C. auratus s.s. They consider C. gibelio to be a distinct species as long as it has a unique and ancient origin rather than arising de novo, and as long as the type specimens are triploids. Szczerbowski in Bănărescu and Paepke (2002) recognises C. a. auratus and C. a. gibelio. Kalous et al. (2012) refer to C. carassius from most of Europe and western Siberia, C. gibelio in Europe, Siberia and northeast Asia with C. auratus in mainland East Asia (and presumably widely introduced). Diploid and triploid individuals of C. gibelio occur in many populations, complicating species definition. Using cytochrome b, Kalous et al. (2012) demonstrated that C. gibelio may comprise two species under that name.

Additionally C. auratus may be a tetraploid derivative of Carassius carassius. The native distribution of C. carassius is in Europe and western Asia, reaching northern drainages of the Caspian Sea in the southern limits of its distribution (Libosvárský, 1962). It differs from C. auratus in having a slightly convex margin to the dorsal fin (straight or slightly concave in C. auratus), caudal fin slightly emarginate (deeply emarginate), usually 6 branched anal rays (always 5), 23-33 gill rakers (37-53), 31-34 vertebrae, usually 32-33 (28-31, usually 29-30), 28-29 fin denticles posteriorly on the dorsal fin spine (10-11), peritoneum light (dark), black spot at the caudal fin base in young and some adults (absent), and a coppery gold body (silvery, pinkish gold, gold or red) (Szczerbowski in Bănărescu and Paepke, 2002). Berg (1948-1949) also cites the characters body rounded, back thick (body angular, back compressed) and scales weakly sculptured (rough), although his comparison is with C. a. gibelio.

Goldfish commonly hybridise with Cyprinus carpio to further confuse the identity of these fishes (L. Nico, http://nas.er.usgs.gov/fishes/accounts/cyprinid/ca_aurat.html, downloaded 24 May 2000). The identity of "goldfish" in Iran has not been thoroughly surveyed and, along with conflicting views on species and widespread introductions from many sources, make it simpler to refer to this taxon as C. auratus for now.

Al-Mukhtar and Al-Hassan (1999) describe a hybrid of this species and Barbus (= Mesopotamichthys) sharpeyi from Al-Hayei (= Al Ha'i), a seasonal lake between the Karkheh and Dez rivers in Khuzestan. Jawad et al. (2012) report C. gibelio from Basrah Province of southern Iraq, distinguishing it from C. auratus and C. carassius.

Key characters

The combination of spines in both the dorsal and anal fins and the absence of barbels is unique to this species. Szczerbowski in Bănărescu and Paepke (2002) distinguishes the subspecies auratus from gibelio by 21-36 lateral line scales (27-35 in gibelio) and a pink or gold colour (yellowish silver), not very diagnostic. Ilhan et al. (2005) give gill raker numbers of 34-40 for auratus, 42-56 for gibelio and 25-32 for C. carassius in Turkish waters (however note below that counts can increase with growth and see also under C. carassius for somewhat different counts and other distinguishing characters).

Morphology

Dorsal fin with 3-4 unbranched rays followed by 12-20 branched rays, anal fin with 2-4, usually 3, unbranched rays followed by 5-6, usually 5, branched rays, pectoral fin branched rays 11-18, and pelvic fin branched rays 6-9, usually 8. Dorsal and anal fin spine denticles coarse and few (about 10-15).

Lateral line scales 21-36. The anterior scale margin is wavy and there are very few anterior and posterior radii, as few as 3-4. The focus is slightly subcentral posterior. Circuli on the exposed part of the scale are more coarse and widely spaced than on the concealed part of the scale. Gill rakers long with serrated interior margins, reaching the fifth to eighth raker below when appressed with younger fish having longer rakers proportionately. Counts are size dependent in the range 34-54. Total vertebrae 25-34. Pharyngeal teeth 4-4, with very elongate, narrow, flattened and horizontal cusps arising from a much narrower stem. The gut is coiled with several loops. This species is variously reported as only diploid or as a tetraploid (2n=100-104); see above.

There are elongate specimens (morpha humilis, where fish density is high) and deep-bodied specimens (morpha vovki, where fish density is low) but these names have no taxonomic significance.

Meristic values for Iranian specimens are:- dorsal fin branched rays 16(4), 17(3), 18(3), 19(5) or 20(2); anal fin branched rays 5(17); pectoral fin branched rays 11(1), 14(1), 15(4), 16(10) or 17(1); pelvic fin branched rays 7(2) or 8(15); lateral line scales 28(6), 29(9) or 30(2); pharyngeal teeth 4-4(10); and total vertebrae 32(2).

Sexual dimorphism

Breeding males have small nuptial tubercles on the operculum, back and pectoral fin rays.

Colour

The golden or orange colour of artificially bred aquarium goldfish is distinctive. However populations in the wild, if they breed successfully, gradually revert to a wild-type of colour, without the appropriate diet supplement of aquarium fish and, as golden fish, are readily seen and eaten by birds and other fishes. Yanar and Tekelioğlu (1999) found that pigmentation increased with fish weight when specimens were fed the carotenoid zeaxanthin. Wild-type colour is an overall olive-green fading to a white belly. Flanks can be silvery to almost black. Fins are a dark olive-bronze, the membranes in particular being heavily pigmented. Young goldfish are usually green, brown or bronze to almost black and only after about 1 year do they take on the colour of adult auratus or gibelio. Peritoneum dusky to black.

Young fish at Ahvaz, Khuzestan, however, are a bright silvery overall (more so than Cyprinus carpio of similar size), the back is grey, the caudal fin is grey on the proximal half and hyaline distally, and the anal fin rays are white (and thus partly resemble gibelio).

Prussian carp (subspecies gibelio) is a dark steel colour with dark blue or greenish dorsally, silver-grey laterally and white ventrally, dorsal and caudal fins are dark grey and the paired fins and anal fin are light pinkish (Szczerbowski in Bănărescu and Paepke (2002).

Size

Attains 62.0 cm and about 5.0 kg, the subspecies gibelio being smaller, up to 45.0 cm and 1.24 kg.

Distribution

The native distribution is in northern Asia and China, reaching northern drainages of the Caspian Sea in the western limits of its distribution (Libosvárský, 1962; Plez, 1987). The goldfish has been widely introduced to garden ponds and released from aquaria in temperate to warm waters world-wide. In Iran it has been introduced throughout the Caspian Sea basin where it is reported from the Atrak, Gorgan, Gharasu, Tajan, Babol, Haraz, Sardab, Aras (including the middle Aras and lower reaches of its tributary the Qareh Chai), Tonekabon, Pol-e Rud, and Safid rivers, the Anzali Mordab where it is now the most abundant fish, Gorgan Bay, Boojagh Wetland and Alma- and Ala-Gol (Holčík and Oláh, 1992; Shamsi et al., 1997; Roshan Tabari, 1997; Abbasi et al., 1999; Kiabi et al., 1999; Abdoli, 2000; Gasmi and Mirzaei, 2004; Patimar, 2008; Abdoli and Naderi, 2009; Khara et al., 2011; Rasouli et al., 2012); the lower Talkheh and lower Zarrineh rivers, teyh Nazlu and Barandoz rivers in the Orumiyeh basin (Abdoli, 2000; Rasouli et al., 2012); the lower Shur, lower Qareh Chai and the Latian Reservoir in the Namak Lake basin (Armantrout, 1980; Hosseini, 1987; Abdoli, 2000); the Hamun Kushk and the Sistan Dam as well as throughout the hamuns in the Sistan basin (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996; field work in the 1970s); Kerman-Na'in and Dasht-e Lut basins generally (Abdoli, 2000); lower Kashaf River in the Tedzhen basin (Abdoli, 2000); throughout Khuzestan where now common (N. Najafpour and M. Al-Mukhtar, pers. comm., 1995; field work 2000, absent in 1970s); in the Qarasu and Yavari spring, Kermanshah (Eagderi and Nasri, 2012); middle and lower Hilleh and lower Mand rivers in the Gulf basin; middle Halil and middle to lower Bampur River (Abdoli, 2000); Dalaki and Shapour rivers (Pazira et al., 2005), Safid River, Zayandeh River, Zarivar Lake and the Hamun Lake (Shamsi et al., 2009), Zarrineh River basin and Zarivar Lake (as C. gibelio; Esmaeili and Gholamifard, 2011), and found in garden and park ponds throughout Iran. Some introductions are probably discarded aquarium fish as goldfish are sold as pets and for the Now Ruz (= New Year) festivities. They may also have been introduced accidentally with the commercially important Chinese carps.

This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters from this source in the Tedzhen (= Hari) River basin.

The Prussian carp (subspecies gibelio) is less widely distributed and its presence and distribution in Iran are not known.

Zoogeography

This species has been introduced to Iran by man. Some are undoubtedly aquaculture pond escapees or aquarium releases. Goldfish are kept in aquaria as part of the Now Ruz (New Year) celebrations in March each year. Tehran television (and the Green Front of Iran, see below) urged people to release them into local waters rather than killing them after the New Year (J. Valiallahi, pers. comm., 2000).

Habitat

Goldfish are hardy and can live in winterkill water bodies with much aquatic vegetation, low oxygen, and high pollution (Gudkov, 1985). They can also survive several hours out of water (Pelz, 1987) and may bury themselves in mud, albeit temporarily when scared (Szczerbowski in Bănărescu and Paepke, 2002). Goldfish appear to favour ponds or pools in streams with aquatic vegetation but are often introduced into small bodies of water as ornamental fish. They are tolerant of turbidity, e.g. clay at 225,000 mg/l, pH from 4.5 to 10.5, very high temperatures (upper lethal limit 41.4°C), and high salinity (17‰). This species was killed under experimental conditions, when gradually acclimated to increasing salinity at 28,200 μmho and, by sudden exposure, at 19,200 μmho (Jassim, 1988). This is a greater tolerance than that shown by Cyprinus carpio, another exotic introduced to Iran. However, Carassius auratus appeared in the Basrah fish market when an increase in the Tigris River discharge reduced the salinity of the Shatt al Arab (N. A. Hussain, in litt., 1994).

In Iran it is one of two most abundant species in Caspian wetland areas along with Gambusia holbrooki (Iranian Fisheries Research and Training Organization Newsletter, 19:4, 1998). Abbasi et al. (2009) in their study of wetlands in Hamadan Province found this species was the second most dominant out of 23 species at 12.5%.

Age and growth

Maturity is attained at 3-4 years in the Volga Delta for goldfish. Life span is 13 years with most growth in the first 2-4 years to a size of 15-20 cm (Gudkov, 1985; Kizina, 1986). Life span in captivity in China may exceed 50 years. Population numbers in confined areas are limited by a chemical released by the goldfish which represses more spawning. Prussian carp live up to 11 years.

In the Anzali Mordab, Holčík and Oláh (1992) found only 6 age groups (as did Bagirova et al. (1990) in reservoirs of Azerbaijan while Pipoyan and Rukhkyan (1998) found 9 age groups in Armenia) with the largest fish 32 cm standard length owing to intense fishing pressure. Growth in mm increments was successively 93, 47, 50, 42, 28, and 37. The population is entirely female (see below). Individual life span is greater in Armenia where males are scarce or absent than in bisexual populations (Pipoyan and Rukhkyan, 1998). Sayad Borani et al. (2001) studied this species (as C. auratus gibelio) in the Anzali Mordab at four localities and found a mean fork length of 19.5 cm (range 2.5-31.5 cm) and a mean weight of 196.8 g. The mean age was 2.6 years. The mean length, weight and age were higher in the Sia-Keshim area of the lagoon. The exploitation rate was 0.47, L was 36.0 cm and K was 0.23 per year. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 41 Iranian fish measuring 5.65-8.17 cm standard length. The a-value was 0.0419 and the b-value 2.911 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases). Patimar (2009) examined fish from the Alma-Gola nd Ala-Gol wetlands in Golestan from 200 to 2002. Ages ranged from 0+ to 8+ with negative allometric growth in Alma-Gol, and positive allometric growth in Ala-Gol. The von Bertalanffy growth curves for mean total lengths were Ltmales = 183.33(1-e-0.31(t+1.05)) and Ltfemales = 245.66(1-e-0.19(t+1.21)) for Alma-Gol and  Ltmales = 224.79(1-e-0.24(t+0.83)) and Ltfemales = 242.80(1-e-0.23(t+0.80)) for Ala-Gol. The sex ratio was unbalanced for males:females at 1:10 and 1:12.7 for Alma-Gol and Ala-Gol respectively because of gynogenesis. Bagheri et al. (2010) examined fish from the Gorgan River estuary and found age groups 1+ to 3+ years, the moment growth coefficient between age groups was 0.57 and 0.32 respectively, and growth was positively allometric.

Fish in Buldan Dam Lake, Gediz River basin, Turkey referred to C. gibelio had a maximum age of 6 years and attained 25.5 cm and 269.1 g (Sarı et al., 2008). von Bertalanffy growth parameters were L = 31.66 cm, W = 635.91 g, k = 0.146 year-1 and t0 = -2.166 year. Ratios of total, natural and fishing mortality were calculated as 0.632 year-1, 0.456 year-1 and 0.176 year-1.

Food

Food is predominately zooplankton but also includes aquatic insects, crustaceans, molluscs, worms, detritus, filamentous algae, macrophytes and young fish, switching from one kind of food to another as circumstances warrant. Goldfish have a palatal organ on the roof of the mouth used to taste and touch food and their dense gill rakers aids in feeding on smaller food items. In the recovering Hawr al Hammar, Iraq, diet is 46.1% algae and 25.5% diatoms, with amounts of plants, crustaceans, insects, snails and fish being less than 10% each, in the Hawr al Hawizah 36.3% algae, 21.3% diatoms and 17.5% copepods, with amounts of plants, cladocerans, ostracods and insects being less than 10% each, in the Al Kaba'ish (= Chabaish) Marsh 45.5% algae, 25.2% diatoms, with plants, various crustaceans, insects and snails at less than 10% each (Hussain et al., 2006).

Reproduction

The fish in the Anzali Mordab are all female, reproducing through gynogenesis. Sayad Borani et al. (2001) found fish in Anzali Mordab to have a sex ratio of 99.3 females:0.7 males. Egg development is stimulated by sperm probably from Cyprinus carpio, Tinca tinca, Blicca bjoerkna or Scardinius erythrophthalmus. Here fish may mature at 1 year of age, and coupled with polycyclic ripening of eggs and intermittent spawning, this has led to the dominance of this species in the fresh waters of the lagoon (Holčík and Oláh, 1992). In Armenia, maturity appears to be linked with average annual temperature - at 12.0-13.1°C it occurs at the end of the first year of life while at 8.4-9.0°C it occurs at the end of third and fourth years (Pipoyan and Rukhkyan, 1998). Turkish populations in Topçam Dam Lake, Aydın (Şaşı, 2008) and Buldan Dam Lake, Gediz River basin (Sarı et al., 2008) referred to C. gibelio were 98.84% and 99.44% female. Spawning in the former locality was from March to August, suggesting multiple spawnings with mean fecundity ranging from 37,823 in August to 85,159 in March. Egg diameter reached 1.099 mm in June.

Patimar (2009) examined fish from the Alma-Gola nd Ala-Gol wetlands and found reproduction in February, March and April. Absolute fecundity reached 13,020 eggs. Gorgan River fish sampled by Bagheri et al. (2010) were all female,

Spawning begins in late April to mid-May in the Volga Delta and occurs in May-June in the Anzali Mordab (Sayad Borani et al., 2001). Eggs are laid in 2-5 batches over a spawning period extending into July. Up to 10 batches are laid elsewhere at 8-10 day intervals with up to 4000 greenish-yellow eggs in each batch. Fecundity reaches 253,200 eggs (elsewhere to 685,700 with absolute fecundity reaching 860,000 eggs). The largest eggs are 1.6 mm in diameter (Gudkov, 1985; Kizina, 1986; Szczerbowski in Bănărescu and Paepke, 2002). Each female is accompanied by 2 or more males and chases are reported with splashing and shooting through the water near the surface. The eggs are adhesive and attach to water plants and hatch in 5-8 days.

Parasites and predators

Mokhayer (1976b) records infectious dropsy and swimbladder inflammation in Iranian goldfish. Saprolegniosis has been reported from goldfish in Iran (Rahbari and Razavilar, 1982). Growths of the fungus Saprolegnia parasitica resembled tufts of cotton wool. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990a) record the monogeneans Dactylogyrus baueri, D. extensus, D. formosus and D. vastator from this species in the Safid Rud. Jalali and Molnár (1990b) report a variety of monogeneans from this species variously in fish farms throughout Iran, namely Dactylogyrus baueri, D. dulkeiti, D. formosus, D. vastator and D. vastator forma minor. Molnár and Jalali (1992) record the monogenean Dactylogyrus intermedius from this species in a petfish farm near Tehran. Gussev et al. (1993a) describe a new species of monogenean from goldfish on a fish farm near Tehran, Dactylogyrus intermedioides. Shamsi et al. (1997) report Clinostomum complanatum, a parasite causing laryngo-pharyngitis in humans, from this species. The helminth Anisakis sp. is recorded from the guts of this species in the Anzali wetland (Ataee and Eslami, 1999; www.mondialvet99.com, downloaded 31 May 2000). Mousavi (2003) records the monogeneans Gyrodactylus sp., G. kabayashi, D. extensus, D. baueri, Trichodina sp., the ciliates Ichthyophthirius multifilis and Ichthyoboda sp. and the copepods Lernaea cyprinacea and Argulus foliaceus from this species in ornamental fish in Iran. Aquarium specimens are often released in the wild at New Year (Now Ruz). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species, monogenean trematodes Dactylogyrus anchoratum, and Gyrodactylus sp.. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum from this species in Lake Zarivar. Naem (2002) records the monogenean Dactylogyrus anchoratus from fish in Safid River. Mehdipoor et al. (2004) record the monogenean Dactylogyrus baueri in this fish in the Zayandeh River. Masoumian et al. (2005) recorded the protozoan parasite Ichthyophthirius multifilis from this species in the Aras Dam in West Azarbayjan (species identified as C. carassius, presumably goldfish). Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. kobayashii and G. sp. in fish from the Safid River. Khara et al. (2006a) record the eye fluke Diplostomum spathaceum for this fish in the Amirkalayeh Wetland in Gilan. Sattari et al. (2004, 2005) surveyed this species (as C. carassius) in the Anzali wetland, recording Raphidascaris acus (and larvae) Eustrongyloides excisus and Camallanus lacustris. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, and found Eustrongylides excisus. Sattari et al. (2007) record the nematode Raphidascaris acus, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus,and Gyrodactylus sp. in this species in the Anzali wetland of the Caspian shore. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Shamsi et al. (2009) found Dactylogyrus baueri, D. dulikeity, D. extensus, D. intermedius, D. intermedioides and D. wegeneri in this species from localities such as fish farms, the Safid River, Zayandeh River, Zarivar Lake and the Hamun Lake. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Lernaea sp. on this species. Khara et al. (2011) list the mongenean Dactylogyrus sp. from this fish in the Boojagh W etland of the Caspian Sea. Mousavi et al. (2011) found that over 19% of goldfish from 10 ornamental fish farms in Iran were infested with Argulus species, and also with Gyrodactylus, Dactylogyrus and Trichodina species and with Ichthyophthirius multifilis. The uncontrolled import of live fish, and their parasites, could cause a serious loss of native fishes. Tavakol et al. (2011) studied occurrence and distribution of the bacteria Aeromonas spp. in farmed goldfish in Gilan, meant for the Now Ruz trade. This bacterium can cause diarrhoea in humans, especially children, and is prevalent in the crowded conditions. Jahanbakhshi et al. (2012) investigated the toxic potential of nanometer-sized particles of silver, used as coatings in various applications and found in ecosystems. Omidzar et al. (2012) identified Gyrodactylus gurleyi on ornamental fish using molecular and morphometric methods. Tarkhani and Imanpoor (2012) evaluated the use of salt and formaldehyde, used against ectoparasites, in terms of stress response, finding a rapid stress which is however eliminated after 24 hours in fresh water. Salt was less stressful. Rasouli et al. (2012) list various ectoparasites from C. carassius (possibly C. auratus) in waters of West Azerbaijan namely Diplostomum spathaceum, Dactylogyrus sp., Gyrodactylus sp., Trichodina sp., Ichthyophthirius multifilis, Argulus sp, and Chilodonella sp.

Ashoori et al. (2012) found that grey herons (Ardea cinerea) in the Siahkeshim Protected Area of the Anzali Wetland ate this species (identified as C. carassius) predominately.

Economic importance

This species is raised on Tehran and Gilan fish farms for the pet trade (Molnár and Jalali, 1992; Tavakol et al., 2011). It forms part of the Now Ruz (= New Year, usually 21 March) celebrations in Iran where a bowl with goldfish forms part of the traditional Haft Sin table setting (so called for seven items that must be present, all beginning with the letter "S", each having a symbolic meaning, the goldfish is in addition to these). The goldfish in a bowl represents life within life, and the sign of Pisces which the sun is leaving.

Haft Sin Table, Iranian Embassy, Ottawa, 2009, photograph by Brian W. Coad
Haft Sin Table, Iranian Embassy, Ottawa, 2009, photograph by Brian W. Coad

Aquarium goldfish at Iranian Embassy, Ottawa, 2009. photograph by Brian w. Coad
Aquarium goldfish at Iranian Embassy, Ottawa, 2009. photograph by Brian w. Coad

Ebrahimzadeh Mousavi and Khosravi (2004) report suspected epizootic ulcerative syndrome from ulcerated goldfish.

In the Anzali Mordab, 62% of the total catch is goldfish, an accidental introduction (Petr, 1987). The catch in the mordab in 1990 was 46,472 kg (Holčík and Oláh, 1992). As the salinity of this lagoon increases, the density of goldfish will decrease. Valeipour and Haghighy (2000) record the catch for 1992-1996 at 40% of the species taken. Safaei (2005) gives a goldfish catch figure of 45% of the 313 ton fishery there in 1992. The presence of goldfish in the Anzali Mordab led to a decline in the native fishery there.

This species is caught by anglers at Ahvaz in Khuzestan using bread or potato as bait.

It is known to control mosquito larvae in Bengal (Chandra et al., 2008).

The peculiar type of reproduction is very successful and affects the catches of other cyprinid species, being equivalent to a predatory effect (Holčík and Oláh, 1992).

There is some evidence that this fish disturbs the habitat of native species, muddying waters, and it may compete for food and space. Goldfish have destroyed some amphibian populations in other parts of the world by consuming frog eggs (Coad and Abdoli, 1993b). The Green Front of Iran recommended the release into pools of mosques, parks or natural lakes of the estimated 20 million goldfish kept in aquaria for the Iranian New Year celebrations in March each year. This would have a deleterious effect on habitats not yet colonised by this exotic species. A news report in 2005 cites the death of 5 million fish in transit from the store to the Iranian home at New Year, indicating perhaps that the numbers that do make it are much higher (www.politicalgateway.com, downloaded 5 August 2005). Newspaper articles suggested that goldfish should only be released into "pools" rather than rivers because of all the attendant dangers of this exotic. They are known to prevent reproduction of native species in Sistan (Iran Daily, 17 March 2005, p. 5).

This species is used in Iran as an experimental organism and for studies in reproductive biochemistry, e.g. in studying the effects of anionic detergents (shampoos, a common water pollutant) on blood parameters, on hepatic and renal pathology and serum biochemical parameters (Shahsavani et al., 2003; Shahsavani et al., 2004; Shahsavani et al., 2005; Shahsavani and Movassaghi, 2003); the use of phenytoin sodium on skin wounds (5mg/l showed best healing improvement while zinc oxide was not as effective) (Shahsavani et al., 2001, 2002, 2002); the formation of lesions and clinical changes in fish exposed to kerosene (Shahsavani et al., 2003); the effects of cortisol on testicular apoptosis (Bahmani et al., 2007); the adverse effects of phenytoin sodium, a drug used for healing skin lesions, on the gills, liver and kidney (Shahsavani et al., 2007); hormonal GnRHa and pituitary extract proved more effective on spermatological parameters than the hormone HCG (Zadmajid et al., 2008); the effects of seminal plasma indices on sperm motility (Zadmajid and Imanpour, 2009); the effects of hormones on seminal plasma biochemistry (Zadmajid et al., 2009); the effects of ascorbic acid on hatching performance temperature tolerance (as C. gibelio) (Taati et al., 2010); coelomic fluid composition and its effects on sperm motility (as C. gibelio) (Taati et al., 2010a); correlation between chemical composition of seminal plasma and sperm motility (as C. gibelio) (Taati et al., 2010b). Kashani et al. (2012, 2012) have examined the significant effect of dietary vitamin C and E and highly unsaturated fatty acid on blood parameters, gonad characteristics, hatching rate and fertilisation success. Tarkhani et al. (2012) found improved fecundity and gamete quality in fish fed diets with 17•-estradiol, enabling better management of egg production for the Now Ruz culurists.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in textbooks, in aquaria and in aquaculture, as bait, as an experimental species and because it has been introduced outside its natural range. There are numerous, commercial aquarium forms with particular morphologies and colours that are assigned common names, e.g. common, veiltail, comet, fans, calicoe, black-moor, telescope-veiltail, lionhead, egg-fish, shubunkin. Balon (2006) reviews the origin of the species.

Conservation

This species is a successful exotic, in no need of conservation.

Further work

The Carassius species in Iran is generally regarded as C. auratus, the goldfish of aquaria, as it is used extensively in Now Ruz (New Year) celebrations and often released into natural waters. This needs confirmation for all major populations.

Sources

Iranian material: CMNFI 1979-0230, 41, 14.7-38.6 mm standard length, Sistan, Hamun-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1991-0162, 1, 40.5 mm standard length, Mazandaran, Bagher Tangeh (36º42'N, 52º43'E); CMNFI 1993-0136, 64.0 mm standard length, uncatalogued material, 1, 93.5 mm standard length, Gilan, near Hendeh Khaleh (ca. 37º23'N, ca. 49º28'E); 1, 52.8 mm standard length, Gilan, near Hendeh Khaleh (ca. 37º23'N, ca. 49º28'E); 4, 16.4-50.3 mm standard length, Gilan, near Khoshk Bijar (ca. 37º22'N, ca. 49º47'E).

Carassius carassius
(Linnaeus, 1758)

Carassius carassius from Wikimedia Commons.

 

Carassius carassius from Wikimedia Commons
Carassius carassius from Wikimedia Commons

Carassius carassius from Wikimedia Commons
Carassius carassius from Wikimedia Commons

The crucian carp has been reported as introduced to Iran in the Karun River basin as aquarium releases by Armantrout (1980) without further details and there are other reports such as in the Gorgan River (Y. Keivany, in litt., 1992) and Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000) but these may be confusion with Carassius auratus. Specimens are needed to confirm the presence of this species in Iran. The native distribution is in Europe and western Asia, reaching northern drainages of the Caspian Sea in the southern limits of its distribution (Libosvárský, 1962). It differs from C. auratus in having a slightly convex margin to the dorsal fin (straight or slightly concave in C. auratus), caudal fin slightly emarginate (deeply emarginate), usually 6 branched anal rays (always 5), 23-33 gill rakers (37-53), 31-34 vertebrae, usually 32-33 (28-31, usually 29-30), 28-29 fin denticles posteriorly on the dorsal fin spine (10-11), peritoneum light (dark), black spot at the caudal fin base in young and some adults (absent), and a coppery gold body (silvery, pinkish gold, gold or red) (Szczerbowski in Bănărescu and Paepke, 2002). Berg (1948-1949) also cites the characters body rounded, back thick (body angular, back compressed) and scales weakly sculptured (rough), although his comparison is with C. a. gibelio, itself recognised as a distinct Carassius gibelio (Bloch, 1782), also of uncertain occurrence in Iran.

Carassius gibelio
(Bloch, 1782)
 

Carassius gibelio from Wikimedia Commons
Carassius gibelio from Wikimedia Commons

The Prussian carp may occur in Iran (see discussion under Carassius auratus above) but distribution and accounts of biology are confused with the latter species and cannot be separated out here. Taati et al (2010, 2010a, 2010b) consider their experimental material from a hatchery at Nahar Khoran, Gorgan to be Prussian carp.

Genus Chondrostoma
Agassiz, 1832

The nases are found from the Iberian Peninsula and France to the Caspian and Tigris-Euphrates basins. There are about 26 species of which 2 are known for Iran (Elvira, 1997). Chondrochylus Heckel, 1843 and Chondrochilus Heckel, 1843 are synonyms. Eschmeyer (1990) gives the year of publication for the genus as 1832 as opposed to other authors who give 1835 (e.g. Berg, 1948-1949; Reshetnikov et al., 1997). Doadrio and Carmona (2004) confirm the monophyly of the genus based on the cytochrome b gene with vicariant events accounting for distribution of taxa better than a dispersalist model. Middle East taxa belong to a single lineage with the more differentiated and basal species in the Caucasus and Mesopotamia, having been isolated in the Upper Miocene-Pliocene.

This genus is characterised by being of moderate size, with a somewhat compressed body, scales of moderate to small size (44-106 in the lateral line (Robalo et al. (2007) give a range of 52-78 for their more restricted genus)), scales squarish with radii in the anterior and posterior fields and a subcentral anterior focus, no barbels, an inferior and transverse or crescentic mouth with a cutting edge to the lower jaw, thin upper lip and no lower lip, pharyngeal teeth knife-like and in 1 row with a high count (5, 6 or 7, the same number on each arch or one more on the left), gill rakers short and moderately numerous (up to 40), short dorsal fin without a thickened ray opposite the pelvic fins, 7-10 dorsal fin branched rays, a moderately elongate anal fin with 8-12 branched rays, deeply forked caudal fin and usually concave dorsal and anal fins, a pelvic axillary process always present, 42-49 vertebrae, a black peritoneum, and a long, coiled gut. Elvira (1997) and Robalo et al. (2007) give osteological characters.

Bogutskaya (1997a) places the nominal Iranian species, C. regium and C. orientalis, in a group characterised by a straight or only slightly arched mouth cleft, high vertebral counts (total vertebrae modes 45-47 and abdominal modes 26-28) and often or commonly 4 unbranched rays in the dorsal fin.

Chondrostoma cyri
Kessler, 1877


Ventral head


Aras River, March 2012, courtesy of K. Abbasi
 

Common names

shekamsiah-e Aras.

[Kur altagizi in Azerbaijan; chernobryushka or blackbelly, Kurinskii podust or Kura nase, uzkotelii Kurinskii podust, all in Russian; Kura undermouth, Kura nase].

Systematics

Earlier works by Elvira (1986; 1988; 1991) placed this species as a subspecies of C. oxyrhynchum but in Elvira (1997), using the phylogenetic species concept and following the studies of Smirnov (1992), this taxon is recognised as a species. C. oxyrhynchum is then found in more northerly rivers of the western Caspian Sea basin remote from Iranian waters. C. cyri orientalis (Bianco and Banarescu, 1982 is described from Fars (see below under C. regium).

Chondrostoma cyri Kessler, 1877 was described from the Kura River, Tiflis (= Tbilisi), Georgia and Chondrostoma oxyrhynchum from the Kuma River near Georgiyevsk, Russia in the Caspian Sea basin.

Alburnus alasanicus Kamenskii, 1901 described in part from the Alasan, Alazan' or Alazani River, a left bank Kura River tributary in Georgia, Chondrostoma schmidti, Berg, 1910 from the Alazan' River at Naporiri, and Chondrostoma leptosoma Berg, 1914 from the Kars-tchai, a tributary of the Aras River in Turkey, the Aras by Kopri-kei, near Erzurum, Turkey, and the lower Aras at Karadonly and Dzhulfa in the former U.S.S.R., are synonyms. Subspecies are not recognised (Elvira, 1991; 1997). C. leptosoma was founded on an elongate form from the Karasu in the Aras River basin.

Two syntypes of Chondrostoma cyri are also in ZISP (10919) from "Tiflis" collected by Kessler in September 1875. A syntype of Chondrostoma oxyrhynchum is in the Zoological Institute, St. Petersburg (ZISP 2881) from "Fl. Sunsha" collected in 1830 by Ménétries. According to Elvira (1988), the type locality is Kuma R. at Georgijewsk and 2 syntypes are under ZISP 10922. Another syntype of Chondrostoma oxyrhynchum is in the Natural History Museum, London (BM(NH) 1897.7.5:28 (184.8 mm standard length), formerly in ZISP, as is other syntype of Chondrostoma cyri (BM(NH) 1897.7.5:27 (correctly numbered 27, 78.4 mm standard length), formerly in ZISP)(Elvira, 1988; personal observations).

Five syntypes of Chondrostoma leptosoma are in the Zoological Institute, St. Petersburg (ZISP 9098) according to (Elvira, 1988) but there are 15 syntypes under this number from the "Reka Araks", 1888, Warpochowsky as well as additional material listed as syntypes with numbers ZISP 9107 ("Fl. Araxes", 1888, Warpochowsky, 12 fish), ZISP 5180 ("Kars-tschai", 1879, Dr. A. Brandt, 3 fish), ZISP 9099 ("Reka Araks", 1888, Warpochowsky, 4 fish), ZISP 15264 ("Reka Araks", 20.III.1911, 2 fish), and ZISP 15516 ("Reka Araks near settlement Djulfa", 17.VI.1911, 13 fish).

Key characters

This species is the only one in its genus in northern Iran and can be recognised by generic characters.

Morphology

Kuru (1981) gives the following meristic characters for 103 specimens from the Aras and Kura river basins in Turkey:- 10-12 dorsal fin rays, 10-11 anal fin rays, 9-10 pelvic fin rays, 9-15 pectoral fin rays, 52-62 lateral line scales, 13-18 scales around the caudal peduncle, 17-32 gill rakers, and 5-6 pharyngeal teeth on each arch (note that the statistical treatment in this paper is in error and the conclusion that species of Chondrostoma in Turkey are not distinct is therefore incorrect). There is clinal variation in scale numbers, the number increasing from south to north and Elvira (1988; 1991) gives the total range for characters of this species as dorsal fin branched rays 7-9, usually 8, anal fin branched rays 8-10, usually 9-10, pectoral fin branched rays 13-18, usually 14-16, pelvic fin branched rays 7-8, usually 8, lateral line scales 50-68 (to 73 in Kazancheev (1981) and from 48 in Chikova (1967)), scales above the lateral line 7-10, usually 8-10, scales below the lateral line 3-6, usually 4-6, pharyngeal teeth 6-5 or 5-5, more rarely 6-6 and mode 6-5, and gill rakers 21-29. Vertebrae number 43-45.

The mouth is arched with a thin horny layer on the lower jaw. Scales are rounded in overall shape with indentations above and below a central, rounded protuberance on the anterior margin. The anterior margin may be wavy. There are few anterior and posterior radii, few circuli and a subcentral anterior focus. There is a pelvic axillary scale. The gill rakers are short and reach the one below or just past it when appressed. Pharyngeal teeth are compressed and thin but deep with a long, thin and concave grinding surface. Teeth tips may be slightly hooked. The gut has numerous anterior loops.

Thirteen specimens from Djulfa (presumably in Azerbaijan opposite the Iranian town across the Aras River) have dorsal fin branched rays 8(12) or 9(1), anal fin branched rays 9(9) or 10(4), and pharyngeal teeth 6-5(5) or 6-6(1).

Sexual dimorphism

Unknown.

Colour

The flanks are silvery but may have dark pigment spots which may, or may not, form a stripe. Paired fins are orange to reddish and median fins grey. The dorsal and caudal fins have dark margins. The peritoneum is black.

Size

Reaches 80.0 cm and about 5.0 kg.

Distribution

Found in the rivers draining to the western coast of the Caspian Sea from the Kuma River in the north southward to the Kura and Aras river basins in the south. Recorded from the Aras River basin of Iran (Abdoli, 2000).

Zoogeography

This genus has a European and Middle Eastern distribution. Its relationships to other taxa are poorly known.

Habitat

Unknown. Found principally in streams and rivers.

Age and growth

Fish are mature at 2 years of age and life span is at least 5 years.

Food

Diet is assumed to consist of bottom organisms including aquatic insect larvae, detritus and vegetation scraped from the substrate.

Reproduction

Up to 16,217 eggs are produced and maximum diameter is 1.69 mm. The spawning season is in the spring, peaking in April in the Kura River basin (Abdurakhmanov, 1962).

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

Kiabi et al. (1999) consider this species to be conservation dependent, in the south Caspian Sea basin according to IUCN criteria. Criteria include sport fishing, possibly few in numbers, limited range (less than 25% of water bodies), absent in other water bodies in Iran, absent outside the Caspian Sea basin.

Further work

Biology in Iranian waters needs study.

Sources

Morphology based on Bianco and Banarescu (1982), Elvira (1986; 1988), Nelva et al. (1988).

Type material: See above, Chondrostoma cyri (BM(NH) 1897.7.5:25, formerly in ZISP), Chondrostoma oxyrhynchum (BM(NH) 1897.7.5:28, formerly in ZISP), and Chondrostoma leptosoma (ZISP 15516).

Iranian material: None.

Comparative material: CMNFI 1980-0812, 2, 101.9-107.9 mm standard length, Turkey, Kars, Selim Çayi (40º28'N, 42º47'E).

Chondrostoma orientale
Bianco and Banarescu, 1982

Chondrostoma cyri orientalis Bianco and Banarescu, 1982 was originally described from the "Pulwar River near Persepolis".

The holotype (IZA 8170, 93.7 mm standard length, examined by me) and 19 paratypes (IZA 7833, 51 specimens under this number, 35.4-90.1 mm standard length) of Chondrostoma cyri orientalis are in the Istituto di Zoologia dell'Universitá di L'Aquila, Italy (Elvira, 1988). Two paratypes of Chondrostoma cyri orientalis are stored in the Field Museum of Natural History, Chicago (FMNH 94519)(Ibarra and Stewart, 1987), 1 paratype is in the Muséum national d'Histoire naturelle, Paris (1982-1014), 1 paratype is in the United States National Museum, Washington (USNM 227934), 2 paratypes are in the Academy of Natural Sciences, Philadelphia (ANSP 150985), and 6 paratypes are in the Canadian Museum of Nature, Ottawa (CMNFI 1982-0365, formerly IZA 7833, 37.8-88.7 mm standard length). The total number of paratypes is 75, originally under IZA 7833 but some dispersed as noted above, with 10 further fish in the Institutul de Stiinte Biologice, Bucurešti, Romania (ISBB) but uncatalogued (Bianco and Banarescu, 1982).

Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found  Lernaea cyprinacea on this species (as C. regium).

 

Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Unio sp., Lernaea cyprinacea, Ichthyophthirius multifilis and Diplostomum spathaceum.

Chondrostoma regium
(Heckel, 1843)

Haramabad, Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Haramabad, Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi

Common names

jokhorak, nazok, nazi; heif-e nan (= waste of bread, i.e. valueless) in Khuzestan; سياه ديم (= siah deem in Behbehan); سياه دم (= siah dom, meaning blacktail); كپور پوزه دار (= kapur puzeh dar).

[baloot muluki, pangki; zurri (= the harmful one) at Mosul (also used for Alburnus mossulensis, Aphanius spp., Gambusia and any small fishes or large fishes when young); terris or terris achmar meleki (= royal red terris) at Aleppo (= Haleb, Syria), all in Arabic; based on Heckel (1843b) for zurri and terris; king nase].

Systematics

Chondrochilus regius Heckel, 1843 was described from the "Orontes" (= Asi) (but see below) and "Tigris". Elvira (1988; 1991; 1997) considers Chondrostoma orientale to be a valid species while Nelva et al. (1988) retain it as a subspecies of C. cyri. Bianco and Banarescu (1982) placed orientalis in C. cyri on the basis of similar dorsal and anal fin ray counts, scale counts and to a certain degree pharyngeal tooth formula.

Banarescu (1960) regarded C. regium as only a race of a widespread species, C. nasus (Linnaeus, 1758). C. nasus has larger scales on average and 6-6 pharyngeal teeth (Berg, 1949); Heckel (1847c) found 47 C. nasus from the Danube River had 6-6 teeth, 2 had 6-7 and 2 had 5-6 while in 13 C. regium the count was 7-6 in 12 fish and 6-6 in 1 fish. Krupp (1985c) considers C. regium to be distinct while recognising the small degree of morphological variation between species in this genus. Data gathered for Iran show a wide range in scale and teeth counts (see below). Ladiges (1960) identified specimens from the same bodies of water in Turkey as members of both species. The earlier literature on the systematics of this genus remain confused (see Elvira (1988) for comments on Ladiges (1966) and Kuru (1981)) and the morphology summarised here for this species does not adequately resolve the problem. There may well be significant variation of a clinal nature, altitude and temperature may be important, and habitat types (lentic or lotic) may affect body form. Most samples examined previously are too small in numbers and differences due to size and sex could not be adequately assessed.

Twelve syntypes of Chondrostoma regium are in the Naturhistorisches Museum Wien (7 fish as NMW 52532-52535 from the Quwayq (= Kueik) River near Aleppo and 5 fish as NMW 52536-52538 from the Tigris River near Mosul)(Elvira, 1988). Krupp (1985c) gives further details. All material was collected by Th. Kotschy in 1842 from the Quwayq and in 1843 from Mosul and the range in standard length for the fish from the Quwayq is 102-166 mm and from Mosul 11.9-24.1 cm. The Vienna catalogue lists only 6 fish but the card catalogue in 1997 lists NMW 52532 (2 fish), 52533 (2), 52534 (2), 52535 (1), 52536 (2), 52537 (1) and 52538 (2) as syntypes. The type locality "Orontes" (= Asi) in Heckel (1843b) seems to be an error.

Key characters

This species is the only one in its genus in southern Iran and can be recognised by generic characters.

Morphology

?re-work

The following counts are from literature sources; my counts in the table below often show a wider range: lateral line scales 56-72 (47-55 for orientalis), scales above the lateral line 9-13 (8-9 in orientalis), and scales below the lateral line 5-6 (4-5 in orientalis). Lateral line scale counts for Iranian fish are as follows: Tigris - 50(1), 51(4), 52(3), 53(12), 54(5), 55(7), 56(4), 57(4), 58(4), 59(8), 60(7), 61(6), 62(5), 63(3), 64(3), 65(1), 66(2), 67(1) or 69(2); Kor (= orientalis) - 49(1), 50(2), 51(3), 52(8), 53(3), 54(7), 55(3) or 57(4). Despite a lower range, the counts for the Kor River basin are matched by a sample from Cheshmeh Javari near Ravansar, Kermanshahan (CMNFI 1979-0287) which have a range of 50-58, leading to a supposition of altitudinal or habitat variation :-

Dorsal fin branched rays 8-11, mode 9 (note Bogutskaya (1997a) gives a mode of 10) (7-9, mostly 8 for orientalis), anal fin branched rays 9-12, mode 11 (note that Bogutskaya (1997a) gives modes of 11 or 12) (9-10, mode 9 for orientalis), pectoral fin rays 14-18, mostly 15-17 (13-15, mostly 14 in orientalis) and pelvic fin rays 6-9, mostly 8 (7-8, mostly 8 in orientalis). Gill rakers 18-36 (probably some lower counts are of rakers on the lower arch only and ranges in single studies, presumably to a consistent technique, are 22-34, 24-31, 25-34 and 25-36) (22-28, 22-30 or 28-32 by different authors for orientalis). Counts for the whole arch on Iranian fish give a wide range of 19-34, highly correlated with size, larger fish having more (or more discernible) rakers than smaller fish (r = 0.5049, p<0.001, n = 90).

Scale radii are few and restricted to the posterior field. Total vertebrae 46-48. Pharyngeal teeth 6-5, 6-6, 6-7, 7-5, 7-6 and 7-7, mode 6-6 or 7-6 (6-5, 6-6, 5-6 and 7-5, mostly 6-6 for orientalis) but see above. The mouth is straight (= transverse) with a thick horny layer on the lower jaw. Esmaeili et al. (2010) give a diploid chromosome number of 2n=52 with 21 pairs of submetacentric and 5 pairs of subtelocentric chromosomes from the Fahlian River in Fars. The arm number was 58.Other Chondrostoma species have 2n=50.

Meristics for southern Iranian specimens of Chondrostoma:


Locality/Dorsal Fin Branched Rays

7

8

9

10

x

S.D.

Tigris River Basin

35

46

1

8.6

0.520

Kor River Basin

1

30

8.0

0.180

Locality/Anal Fin Branched Rays

8

9

10

11

12

x

S.D.

Tigris River Basin

4

45

21

9

3

9.5

0.892

Kor River Basin

23

8

9.3

0.445

Locality/Pelvic Fin Rays

7

8

9

x

S.D.

Tigris River Basin

3

77

2

8.0

0.248

Kor River Basin

1

29

1

8.0

0.258

Locality/Lateral Line Scales

Range

x

S.D.

Tigris River Basin

50-69

57.8

4.553

Kor River Basin

49-57

53.2

2.131

 

Locality/Total Vertebrae

42

43

44

45

46

47

48

49

x

S.D.

Tigris River Basin

1

20

17

9

14

10

9

2

45.1

1.833

Kor River Basin

5

20

5

1

43.1

0.680

Sexual dimorphism

Unknown.

Colour

The back is olive-brown with bluish tinges and the flanks and belly are silvery-white. The dorsal and caudal fins are greyish and the other fins hyaline. Some fish have bright orange fins, the pectorals paler, the pelvics and anal fins fringed by white. The dorsal and caudal fins have a black margin, wide on the caudal. These fin colours give them a flag-like effect (Heckel, 1843b). The caudal fin can be orange, distally black, with the extreme edge white in freshly dead fish.

Size

Attains 40 cm and 1 kg.

Distribution

Found in the Tigris-Euphrates basin and the Mediterranean basins of southeastern Turkey and the northern Levant. In Iran found in the Tigris River basin. Additional localities are springs (sarabs) near Kermanshah, the Marun River, the Hawr al Azim marsh (Wossughi, 1978; Abdoli, 2000) and the Gamasiab (K. Abbasi, see photograph above. Ghorbani Chafi (2000) lists the Bazoft and Kuhrang rivers in the upper Karun River basin and also possibly the Zayandeh River of the Esfahan basin.

Zoogeography

This genus has a European and Middle Eastern distribution. Its relationships to other taxa are poorly known.

Habitat

Found in both rivers and lakes (and reservoirs) but habitat requirements have not been studied in Iran. Ünlü (2006) reports that this species prefers stone grounds and still waters in rivers and lakes in Turkey.

Age and growth

Khalaf et al. (1986) studied this species in the Diyala River, Iraq. Maximum age group is 7+ years, males and females show no difference in weight at the same length and samples from three adjacent areas show no major differences in growth rates. Length-weight relationship was W = 0.0480 L2.49 (n = 255, r = 0.88). Males mature at 15.0 cm and females at 19.0 cm in the Diyala River at Rustamiyah in Iraq (Allouse et al., 1986). A population at Al Kadhmia north of Baghdad in the Tigris River had four age classes dominated by the three year age class, with all fish being sexually mature during the second year. Fish smaller than 15 cm for males and 17 cm for females were immature. The disparity in age structure with the Diyala River population was attributed to pollution in the Diyala (Daoud and Qasim, 1999).

Polat and Gümüş (1995) aged a population of this species in the Bafra Altınkaya Dam lake in Turkey using vertebrae, otoliths, scales, opercle and subopercle. Age reached 5, perhaps 6, years and scales were found to be the best structure to use. Polat et al. (1999) found a similar age range in the Suat Uğurlu Dam, Turkey with annulus (hyaline ring) formation in October to February. Oymak (2000) examined growth characteristics of this species in the Atatürk Dam on the Turkish Euphrates River. Eight age groups were found and age-length and age-weight equations given for females and males were Lt = 38.67[1-e-0.136126(t+3.073799)], Wt = 527.52[1-e-0.136126(t+3.073799)]3.1986 and Lt = 35.01[1-e-0.168137(t+2.754214)], Wt= 724.73[1-e-0.168137(t+2.754214)]3.2779 respectively. The length-weight relationships were obtained as Log W = -5.4153 + 3.1986 Log FL in females and Log W = -5.6212 + 3.2779 Log FL in males. The condition factor was high in age group 7 and high in April and May, lowest in December and January. Gümüş et al. (2002) found deposition of hyaline rings was synchronous with decrease in food diversity in autumn in the Suat Uğurlu Dam, Turkey. Aydin et al. (2004) demonstrated a positive linear relationship between otolith length and fish length for this species in Keban Dam Lake, Turkey.

Food

This species is omnivorous taking insect larvae and eggs and fry of other fishes. Gut contents also include diatoms and algae as well as large quantities of sand. However, Gümüş et al. (2002) examined diet in the Suat Uğurlu Dam, Turkey and found Navicula, Cymbella and Synedra were the most frequently consumed organisms. This species feeds mostly on Bacillariophyta in this dam but also Chlorophyta, Cyanophyta, Xanthophyta, Euglenophyta and Rotifera. Diet varied with seasonable availability of food items.

Reproduction

Studies on the Diyala River population in Iraq found fish to be mature in December and by January females lacked eggs. Each female produces up to 6900 eggs and number of eggs increases linearly with length (Allouse et al. (1986). The breeding season at Al Kadhmia in the Tigris River near Baghdad was March-May (Daoud and Qasim, 1999). Al-Rudainy (2008) gives sexual maturity as 3 years, 25 cm total length and 250 g weight with spawning in February and March on gravel beds in shallow water with strong current. for Iraq. Ünlü (2006) reports up to 13,280 eggs for fish in the Tigris River of Turkey. Beckman (1962) states that this species probably spawns in May or June in Syria and Oymak (2000) found that condition factors were highest in April and May in the Atatürk Dam, Turkey.

Parasites and predators

Barzegar et al. (2004) examined this species for parasites in fish from the Beheshtabad river in Chahar Mahall va Bakhtiari Province and found Lernaea cyprinacea, Dactylogyrus ergensi, Ichthyophthirius multifilis and Myxobolus sp. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. sp. from the Dez and Karun rivers in Chondrostoma nasus, presumably this species. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Riassy et al. (2009) found the digenean eye parasite Tylodelphys clavata in fish from Choghakhor Lagoon.

Economic importance

This species has been caught and used for food in Khuzestan.

Conservation

This species is relatively common and is not widely used as food; it may not need conservation. However it is listed as endangered in Turkey (Fricke et al., 2007).

Further work

Its biology in Iran needs study and this may reveal conservation needs.

Sources

?re-work

Type material: See above, Chondrostoma cyri orientalis (IZA 8170, IZA 7833, CMNFI 1982-0365, formerly IZA 7833).

Iranian material: CMNFI 1979-0025, 16, 22.1-119.0 mm standard length, Fars, Kor River at Marv Dasht (29º51'N, 52º46'30"E); CMNFI 1979-0028, 14, 32.2-139.1 mm standard length, Fars, Kor River drainage (no other locality data); CMNFI 1979-0059, 1, 72.2 mm standard length, Fars, Pulvar River (30º01'30"N, 52º57'E); CMNFI 1979-0061, 14, 9.5-56.5 mm standard length, Fars, stream tributary to Pulvar River (30º04'N, 53º01'E); CMNFI 1979-0245, 5, 35.3-47.1 mm standard length, Sharestan-e Bahktiari va Chahar Mahall, stream in Ab-e Shalamzar drainage (32º08'N, 50º51'E); CMNFI 1979-0247A, 4, 57.2-65.3 mm standard length, Sharestan-e Bakhtiari va Chahar Mahall (31º57'N, 51º01'E); CMNFI 1979-0248, 2, 39.2-65.2 mm standard length, Sharestan-e Bakhtiari va Chahar Mahall, stream 3 km east of Boldaji (31º55'N, 51º05'E); CMNFI 1979-0271, 11, 60.0-131.3 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0272, 1, 58.5 mm standard length, Lorestan, river at Nokhor (ca. 33º40-47'N, ca. 48º28-45'E); CMNFI 1979-0279, 2, 61.8-134.0 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0280, 1, 114.5 mm standard length, Lorestan, Kashkan River drainage (ca. 33º43-47'N, 48º12-15'E); CMNFI 1979-0283, 1, 137.0 mm standard length, Kermanshahan, river 15 km before Kermanshah (34º21'N, 47º07'E); CMNFI 1979-0287, 22, 56.6-112.5 mm standard length, Kermanshahan, Chashmeh Javari near Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0286, 11, 77.4-100.4 mm standard length, Kermanshahan, Ravansar River at Ravansar (34º43'N, 46º40'E); CMNFI 1979-0289, 1, 131.5 mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0368, 4, 54.0-84.5 mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0370, 6, 187.3-221.6 mm standard length, Khuzestan, Karkheh River (32º12'N, 48º14'30"E); CMNFI 1979-0382, 2, 37.7-62.5 mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0392, 1, 53.7 mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0421, 5, 114.0-122.0 mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Khersan River drainage (30º24'N, 51º47'E); CMNFI 1979-0499, 1, 113.0 mm standard length, Fars, irrigation ditch 32 km from Kor River bridge (30º04'30"N, 52º36'E); CMNFI 1979-0500, 7, 94.8-110.5 mm standard length, Fars, Pulvar River at Naqsh-e Rostam (29º59'N, 52º54'E); CMNFI 2007-0100, 2, 165.4-165.7 mm standard length, Azarbayjan-e Bakhtari, Kalwi Chay near Piranshar (ca. 36º44'N, ca. 45º10'E); CMNFI 2007-0111, 2, 183.3-191.7 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0113, 2, 106.7-145.0 mm standard length, Kermanshahan, Razavar River, Qareh Su tributary (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0115, 3, 72.5-96.5 mm standard length, Kermanshahan, Qareh Su basin north of Kermanshah (ca, 34º34'N, ca. 46º47'E).

Comparative material: BM(NH) 1931.8.12:1-3, 2, 136.0-172.2 mm standard length, Iraq, near Mosul (36º20'N, 43º08'E); BM(NH) 1971.4.2:6, 1, 147.7 mm standard length, Iraq, River Tigris near Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:81-82, 1, 197.5 mm standard length, Iraq, Great Zab near Eski Kelek and near Bekhne Dam (no other locality data).

Genus Crossocheilus
Kuhl and van Hasselt, 1823

Kottelat (1987) retains the spelling Crossocheilus Kuhl and van Hasselt in van Hasselt, 1823 as first reviser. The name was spelt Crostocheilus early in the text but this has never been used again and Crossocheilus appeared with the description. Crossochilus Günther, 1868 is an incorrect emendation (Eschmeyer, 1990). The correct genus for the Iranian species may be Gonorhynchus McClelland, 1839 based on a molecular analysis by Yang et al. (2012)

The genus is found chiefly in the Oriental Region but extends into Iran with one species. There are about 18 species.

The genus is characterised by an elongate body with a rounded belly; the mouth is inferior and transverse, the crenulated or fringed upper lip being continuous with the snout (not separated by a groove); the lower jaw has a horny covering and behind this are several rows of lobate papillae; 1-2 pairs of barbels; gill membranes attached to isthmus; dorsal and anal fins are short and spineless; the lateral line is complete; scales are large to moderate in size; the intestine is very long; and the peritoneum is black.

The lower surface of the head bears an "adhesive apparatus", the mechanism of which has been investigated by Singh (1993) for Crossocheilus latius latius, a subspecies not found in Iran. The fringed upper lip acts as a food strainer as well as part of the adhesive apparatus. Both this lip and the area behind the lower lip are heavily tuberculate with glandular openings and irregularly arranged hard ridges. Mucus from the glands in conjunction with the ridges holds the fish to the substrate.

Crossocheilus latius
(Hamilton, 1822)

Zirdan Dam, Kaju River, 26 December 2011, courtesy of Asghar Mobaraki
Zirdan Dam, Kaju River, 26 December 2011, courtesy of Asghar Mobaraki

Zahak, Sistan River, January 2012, courtesy of K. Abbasi
Zahak, Sistan River, January 2012, courtesy of K. Abbasi

Common names

None.

[ispigoar or dogra in Pakistan].

Systematics

Cyprinus latius was described from the Tista River in India/Bangladesh and types are unknown (Eschmeyer et al., 1996).

Bianco and Banarescu (1982) and Bănărescu (1986) consider Discognathus adiscus Annandale, 1919 described from Sistan (type locality given below) to be a synonym of this species which is represented in Sistan and Baluchestan by Crossocheilus latius diplocheilus (Heckel, 1838), originally described from Kashmir with syntypes in the Naturhistorisches Museum Wien under NMW 48820 (7 fish). Bănărescu (1986) cites 1 fish under NMW 48420 as possibly the holotype but this specimen is dated 1839 which is anachronistic. Berg (1949) considers Discognathus adiscus to be a distinct species. I concur with Bianco and Banarescu (1982) and Bănărescu (1986).

Characters advanced by Berg (1949) for separating the two species are number of barbels (4 in adiscus, 2 in latius diplocheilus where mouth angle barbels are absent or rudimentary, not the rostral ones as implied by Bianco and Banarescu (1982)), upper lip fringe (barely developed in adiscus, distinctly developed in latius diplocheilus), the posterior swimbladder (conical in adiscus, elongate cylinder in latius diplocheilus), and papillae on the lower lip and chin (rudimental in adiscus and latius diplocheilus but the latter has almost free lateral edges and an attached posterior end - this condition is not specified for adiscus). Bianco and Banarescu (1982) and Bănărescu (1986) found some latius diplocheilus specimens to have 4 barbels (and this is given too as a character of C. latius latius), and no difference in development of lip papillae in specimens from the Indus River basin (actually my reading of Berg (1949) cited above does not indicate that papillae development differs but that the sucker area has almost free lateral edges and an attached posterior end; this occurs in Sistan fish but not in 4 fish from the Hamun-e Mashkid and Makran basins of Iran which have a fold in the flesh behind the tuberculate area - these latter fish are very small however, 20.8-27.8 mm standard length, and I lack extensive comparative adult material from outside the Sistan basin in Iran and from neighbouring Pakistan to make an adequate analysis of nominal adiscus and latius diplocheilus in this and other characters). My observations of the posterior swimbladder development indicate a great individual variation in form for Sistan fish: the swimbladder may be conical, elongate and tapering, rounded posteriorly, expanded posteriorly, rounded posteriorly after a constriction, or even a narrow elongate cylinder supposedly characteristic of latius. Fringe development of the upper lip is also quite variable and seems to be relatively well-developed in larger Sistan fish.

Karaman (1971) described a new genus, Hemigarra, for Tylognathus elegans Günther, 1868 and Discognathus adiscus Annandale, 1919. He places Crossocheilus adiscus as the Sistan subspecies of his Hemigarra elegans (= Hemigrammocapoeta elegans here, q.v.) which is found in Mesopotamia. Karaman (1971) distinguishes the two subspecies by the former having densely arranged papillae on the chin as opposed to sparse papillae. Bianco and Banarescu (1982) and Bănărescu (1986) state that it is not related to Hemigrammocapoeta elegans but is a typical Crossocheilus species.

The type locality of Discognathus adiscus is Sistan by implication, as no locality is given for the holotype in Annandale (1919b). Menon and Yazdani (1968) concur. Distribution is given as "small watercourses and pools in the plains of Seistan" and "Nasratabad, irrigation channel in Consulate garden; pool in the desert 5 miles south of Nasratabad; pools in stream-bed 12 miles north of Nasratabad; channels in the reed-beds of the Hamun-i-Helmand near Lab-i-Baring, and channel leading out of the Hamun 12 miles east of Lab-i-Baring; small watercourse, Lutak, southern Seistan", and one of these is presumably the type locality.

Twenty syntypes of Discognathus adiscus are in the Zoological Survey of India, Calcutta (ZSI F9758/1) (Menon and Yazdani, 1968). Annandale (1919b) cites ZSI 9763/1 as the holotype catalogue number. Three syntypes are in the Zoological Institute, St. Petersburg (ZISP 25411) from "Nasratabad, Seistan, Indian Museum, Dr. Hora" and measure 38.0-43.4 mm standard length. Two syntypes (listed as cotypes) measuring 44.8-45.5 mm standard length from "Jellalabad" with the annotation "Ind. Mus. Ex. F 9762/1" are in the Natural History Museum, London (BM(NH) 1919.8.16:7-8; the outside has 1919.3.16:7-8, incorrectly).

Key characters

The characters of the genus, particularly in the mouth region, serve to identify the only species in Iran.

Morphology

Four short barbels are present, the rostral ones longer than those at the mouth corner. The upper lip covers the upper jaw, is granular or tuberculate and has a marginal fringe, variably developed and most apparent in larger fish. The lower lip is only apparent at the sides and the exposed lower jaw has a granular or tuberculate pad without a free posterior margin but with almost completely free edges.

Dorsal fin with 2-3 unbranched and 8-9 branched rays, anal fin with 2-3 unbranched and 5 branched rays, pectoral fin branched rays 14-17, and pelvic fin branched rays 7-9. Lateral line scales 33-39. Scales may have short dorsal and ventral projections from the margin at about one-third of the scale length from the posterior edge. There is a pelvic axillary scale. Scales have 9-10 radii on the posterior field and are elongate with a notably anterior focus. Radii in large fish are parallel rather than divergent. The anus is 4-5 scales in advance of the anal fin origin. Gill rakers 17-25, small reaching the adjacent or second raker when appressed. Pharyngeal teeth usually 3,3,5-5,3,3 or 2,4,5-5,4,2, depending on how the crowded teeth are counted; major row teeth are usually 5 but may be 4 or 6, middle row teeth are 3 or 4, and minor row teeth 2 or 3, more rarely 1 (this difficulty in assigning teeth to rows is the reason for omitting frequency distributions below). Supernumerary teeth may be present to further confuse counts. The crown of major row teeth are flattened, the anterior tooth may be rounded and some teeth may have a small hooked tip. The gut is very long and complexly coiled. The chromosome number is probably 2n=48 (Klinkhardt et al., 1995).

Iranian fish from Sistan and Baluchestan have the following meristic characters: dorsal fin branched rays 8(81) or 9(1), anal fin branched rays 5(81), pectoral fin branched rays 14(28), 15(37), 16(16), or 17(1), and pelvic fin branched rays 7(3), 8(76), or 9(3). Lateral line scales 33(1), 34(2), 35(12), 36(31), 37(32) or 38(4). Total gill rakers 20-25, but not countable with great accuracy since the smallest rakers are difficult to detect at the ends of the arch. Total vertebrae 34(6), 35(20), 36(9) or 37(1).

Sexual dimorphism

Unknown.

Colour

The back is bluish-grey in Sistan fish or brownish to greenish with irregular spots in other populations and the belly light pink to yellowish-white or silvery-white. Fins are pink and the dorsal and caudal fins have a grey tinge. The flank has a bluish, mid-lateral stripe in Sistan fish and in preserved ones scattered melanophores, or small blotches of less than scale size, or clumps of melanophores centred on upper flank scales and more dispersed on the lower flank. There is a broad stripe along the back mid-line. Fins in preserved fish from Sistan are mostly immaculate except in the larger fish with some melanophores lining rays basally. The caudal fin is distinctive in larger fish from Sistan in having the rays of the lower half of the fin heavily pigmented while the upper half rays are only lightly pigmented. Peritoneum is dark brown to black.

Size

Attains 14.6 cm although the largest fish recorded from Sistan was 93.2 mm standard length.

Distribution

Found in submontane areas of Afghanistan, Pakistan and India as well as eastern and southeastern Iran. The main areas of distribution are Sistan, the Hamun-e Mashkid basin including the Simish River and coastal streams of Makran from the Jagin to middle and upper Nikshahr rivers and the middle and upper Bahu Kalat River including its Sarbaz River reach. (Nikol'skii, 1899; Annandale, 1919b; Berg, 1949; Bianco and Banarescu, 1982; J. Holčík, in litt., 1996; Abdoli, 2000; K. Abbasi, see photograph above).

Zoogeography

This distribution in Iran marks the western limit for the genus and the relationships of the species lie to the east.

Habitat

Very abundant in small streams, including those with rocky or muddy beds, irrigation ditches, channels in reed beds and pools in Sistan, less common in Baluchestan streams. This species is found in large schools in Sistan in still or slow-flowing water, on the bottom during the day but it may swim at the surface in the evenings. It is common in the smallest permanent water channels but Annandale and Hora (1920) reported it to be in small numbers in the reed beds in winter and these were dead or dying, perhaps because of low oxygen conditions associated with vegetation decay. Large numbers die each year in drying stream beds as salt content increases and the water is fouled by sheep and goats. Tekrival and Rao (1999) report its aquarium preferences as 18-22°C, pH 6.5-7.2, algae as food, not too bright lighting, bottom dwelling with stones, roots and crevices preferred and cave brooding reproduction. Asghar Mobaraki (pers. comm., 8 January 2012) recorded a fish kill of this species on 26 December 2011 at the Zirdan Dam construction site on the Kaju River.

Age and growth

Unknown in detail. Zare et al. (2011) give length-weight values of 0.032 for a (intercept) and 2.455 for b (slope) for fish from the Chahnimeh Reservoir, Sistan, the b value being low..

Food

Diet is algae on muddy bottoms. The type subspecies is a bottom feeding herbivore taking more than 90% plant food such as algae, diatoms and macrophytes as well as detritus (Sharma, 1984; Singh and Bahuguna, 1984). Iranian fish contain detritus and some insect remains, possibly as accidental inclusions.

Reproduction

Iranian adult specimens were caught in May in Sistan and show signs of developing reproductive organs suggestive of summer spawning.

Parasites and predators

Jalali et al. (2000) describe two new species of monogenean, Dactylogyrus faridpaki and D. eslamii, from this species in the Bahu Kalat River of Baluchestan.

Economic importance

This species is of no economic importance although Butt (1995) suggests it could be cultured as food and as a forage fish in Pakistan.

Conservation

This species does not appear to be under any major threat as it can survive drying of the Sistan lakes in small ditches and streams.

Further work

The biology of this species needs investigation as does the taxonomic status of Sistan populations.

Sources

Mirza (1972) for colour.

Type material: See above, Discognathus adiscus (ZISP 25411, BM(NH) 1919.8.16:7-8).

Iranian material: CMNFI 1979-0224, 8, 43.6-55.4 mm standard length, Sistan, effluent of Hirmand River (30º53'30"N, 61º27'E); CMNFI 1979-0226, 277, 29.7-78.8 mm standard length, Sistan, pool near Kuh-e Khajeh (30º57'N, 61º17'E); CMNFI 1979-0227, 4, 37.0-48.9 mm standard length, Sistan, naizar at Kuh-e Khajeh (30º57'N, 61º16'E); CMNFI 1979-0228, 1, 42.9 mm standard length, Sistan, ditch 1 km from Zabol (31º02'30"N, 61º31'E); CMNFI 1979-0229, 5, 52.3-93.2 mm standard length, Sistan, ditch 5 km from Zabol (31º03'N, 61º33'E); CMNFI 1979-0230, 1, 48.3 mm standard length, Sistan, Hamun-e Puzak (ca. 31º15'N, ca. 61º42'E); CMNFI 1979-0232, 9, 44.0-65.9 mm standard length, Sistan, ditch 11 km from Zabol (ca. 30º58'30"N, ca, 61º36'E); CMNFI 1979-0234, 17, 40.4-49.3 mm standard length, Sistan, effluent of Hirmand River (30º54'N, 61º40'E); CMNFI 1979-0318, 2, 24.0-27.8 mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0333, 2, 20.8-21.2 mm standard length, Baluchestan, Mashkid River west of Kuhak (ca. 27º05'N, ca. 63º12'E).

Genus Ctenopharyngodon
Steindachner, 1866

The grass carp genus contains only a single species found in East Asia but widely introduced for food and its ability to digest macrophytes.

This genus is characterised by a rounded body and broad head, the eyes are large and positioned at or above the body axis and often visible from the underside of the head, mouth wide and terminal, no barbels, moderate-sized scales, a complete lateral line, dorsal and anal fins short and lacking spines, branchial membranes attached to the isthmus, short unfused gill rakers, brown to black peritoneum, and pharyngeal teeth in 2 rows with the crowns strongly compressed and serrate and with a longitudinal groove on the grinding surface.

Ctenopharyngodon idella
(Valenciennes, 1844)

U.S. Fish and Wildlife Service
U.S. Fish and Wildlife Service

http://commons.wikimedia.org/wiki/File:Ctenopharyngodon_idella.jpg
Wikimedia Commons
 

Common names

كپور علفخوار (= kapour-e alaf khaar or alaf khoar or kopur 'laf khoar, carp grass-eater or grass-eater), آمور (= amur), سفيد پرورشي (safid parvareshi or mahid safid parvareshi meaning cultured white fish, from a resemblance to mahi safid, i.e. Rutilus frisii kutum).

[grass carp, white amur].

Systematics

Leuciscus idella was originally described from China. A hybrid of this carp and Rutilus frisii has been bred at the Astaneh Ashrafie Fisheries Research Station and named "Samur" (Iranian Fisheries Research and Training Organization Newsletter, 11:6, 1996). See under Rutilus frisii for more information. Yousefian (2011b) compared Mazandaran and Khuzestan stocks morphometrically and meristically and found significant differences in various characters. The Khuzestan fish were more homogenous and the two populations were considered to be well-established strains with a common nacestor.

Key characters

This species is identified by the eyes being low on the side of the head, the anal fin is far back on the body close to the caudal fin, and pharyngeal teeth have large, parallel grooves on the grinding surface.

Morphology

Lateral line scales 34-47. Scales have a wavy anterior edge, central focus and moderate numbers of anterior and posterior radii. Dorsal fin branched rays 6-8, usually 7, after 3 unbranched rays, anal fin branched rays 7-9, usually 8, after 3 unbranched rays, pectoral fin branched rays 13-20 and pelvic fin branched rays 7-8. Gill rakers number 15-18 and touch the adjacent raker when appressed. Vertebrae 40-47. Pharyngeal teeth are 2,5-5,2, 2,4-5,2, 2,4-4,2, or 1,4-5,2 and are obviously serrated with a longitudinal grooves. The gut is long and complexly coiled. The diploid chromosome number is 48, the triploid 72 (Klinkhardt et al., 1995; Nowruzfashkhami et al., no date). Serum immunoglobulins have been characterised by Soltani et al. (2003).

Sexual dimorphism

Nuptial tubercles are evident on the male head, upper caudal peduncle, dorsal and caudal fins and in particular on the pectoral fins, the first ray of which is thickened, while the female has a distended belly and a swollen and pinkish vent.

Colour

The back is dark, olive to greenish-brown, the flanks are silvery but scales are marked with darker pigment on their posterior margin giving the appearance of a row of spots, and the belly is white to cream-yellow. Scale centres may reflect golden or yellowish tints. Upper scales are outlined with dark pigment to give a cross-hatching effect. The fins are grey-green, or grey to black, except the pelvics which resemble the belly colour. Peritoneum brownish black.

Size

Reputed to attain 1.62 m and about 60.0 kg in its native range (Machacek (1983-2012), downloaded 27 July 2012); reports of weights up to 180 kg probably being exaggerations. Reaches 80 cm in the Tadjan River near Sari (A. Abdoli, pers. comm., 1995).

Distribution

The native distribution is in East Asia but it has been introduced to Iranian waters. Also introduced to Afghanistan, Pakistan, and Iraq (Shireman and Smith, 1983).

This species was first introduced in the 1950s according to Armantrout (1980) in the Anzali Mordab for vegetation control, adults surviving to the 1960s but no breeding population was established. Also introduced in 1966 from a hatchery in the Krasnodar region of the former U.S.S.R. and stocked in the Anzali Mordab (Anonymous, 1970b) and in October 1970 50,000 fingerlings from the U.S.S.R. were introduced to the Caspian Sea and Anzali Mordab (Griffiths et al., 1972). Three large fish (80 cm) were caught in January 1971 and believed to be from the October introduction and evidence of good growth although they may have been from an earlier stocking. It is reported from the Siah-Keshim Protected Region of the Anzali Mordab (Riazi, 1996), presumably recently stocked, and is stocked in a variety of reservoirs in the provinces of Gilan and Mazandaran but not as widely as silver carp (Hypophthalmichthys molitrix). It is pen cultured in Gomishan Reservoir, Mazandaran (Madbaygi, 1993b). Grass carp were introduced to Khuzestan in the 1970s to control vegetation in irrigation ditches. In April 1974, 1150 fish were released in the Dez Irrigation Project (Saadati, 1974). It is reported from Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), from the Safid River and Anzali Talab (Abbasi et al., 1999), from Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), the Kor River in Fars (A. Alamdari, in litt., 1997), from the Sistan basin in Hamun Sabari, Hamun Kushk and the canal flowing into Chahnimeh (Ahmadi and Wossughi, 1988; Mansoori, 1994; J. Holčík, in litt., 1996), from the Haft Barm lakes near Shiraz in 1984 although these later disappeared, possibly eaten by introduced Sander lucioperca (Petr, 1987). As escapees from a fish farm, they have been found in Lake Famur. Also recorded from the Gorgan, Tajan and Safid rivers, and the Anzali Mordab (Kiabi et al., 1999), and it is mapped from the Kor, Kerman-Na'in, Hormuz, Dasht-e Lut and Sistan basins without exact localities; the Kashaf River in the Tedzhen River basin; middle reaches of the Atrak River, lower reaches of the Gorgan, Neka, Babol, Heraz and Safid rivers and in the Anzali Mordab, all in the Caspian Sea basin (Abdoli and Naderi, 2009), the middle to lower Talkheh and lower Zarrineh rivers in the Lake Orumiyeh basin, the middle to lower Abhar-Shur and Qom River in the Namak Lake basin, the middle to lower Zayandeh River in the Esfahan basin; the lower Karun and Jarrahi rivers in the Tigris River basin, and the lower Jovein and middle Kal Shur rivers in the Dasht-e Kavir basin (Abdoli, 2000).

It was introduced to the Soviet Caspian Sea in 1970-1974 where small populations became established in the Terek River and the Volga delta and to the Karakum Canal and Kopetdag Reservoir of Turkmenistan near the Iranian border (Baltz, 1991; Shakirova and Sukhanova, 1994; Sal'nikov, 1995; Opuszynski and Shireman, 1995).

Grass carp could establish breeding populations in the large rivers of southern Iran and Iraq if the environment proves favourable and there is enough uninterrupted river flow for eggs to hatch.

Zoogeography

This species is an exotic in Iran and has a native range from the Amur River basin of Siberia south to southern China. It has been widely introduced around the world for vegetation control.

Habitat

The natural habitat is large rivers but this species adapts easily to pond culture. Grass carp can live in the Caspian Sea at salinities of 5-8‰ although a few are found at 10-12‰. They enter rivers to spawn (Abdusamodov, 1986). Temperatures in the range 0-41°C and low oxygen concentrations (0.2 mg/l) are tolerated by this species as is high turbidity. Fry have an upper lethal temperature range of 33-41°C and temperatures greater than 38°C are lethal for adults. pH range is 5.0-9.0. Adults prefer densely vegetated inshore areas with depths of 1-3 m. Adults leave the river after spawning and feed in lakes, reservoirs and on floodplains, returning to the river in autumn to overwinter in deep holes separate from the juveniles. Young hide in vegetation of lakes, reservoirs and floodplains. Juveniles may migrate as much as 1000 km up- or downstream from the original spawning site in their native habitat. Young fish overwinter in deep holes in river beds.

Age and growth

Growth rate in Khuzestan canals was 1.8 g per day while in ponds growth was 6.6 g per day when fed alfalfa during a 5-month growing season from April to September (Saadati, 1974; Behnke, 1975a). Males begin to mature at 4 years and females at 5 years in the Terek River of Dagestan (Abdusamadov, 1986). Maturity is attained at 6-10 years in the Amur River, the native habitat, and as early as 10 months in Malaysia. Life span is over 33 years. Growth rate in this species is perhaps greater than in any other fish. Growth to 1 kg in the first year of life and 2-3 kg per year thereafter in temperate areas is very high; in tropical areas a 20 g fingerling can reach 8.5 kg in 1 year. Rates of 10-22 g per day have been reported in various areas of the world depending on local conditions.

Food

Grass carp are herbivores, except for quite small fish (20 mm total length or less) which consume zooplankton. In Khuzestan, the grass carp prefers to eat Potamogeton spp. and Alisma gramineum to Chara and Cladophora (Saadati, 1974). The grass carp can consume 100-150% of its body weight per day of aquatic vegetation. Peak feeding occurs at 25-30°C but food is taken in the range 15-35°C. Grass carp stocked in the Anzali Mordab and fish farms of the Caspian Sea basin consume fresh Azolla, an introduced fern. Grass carp stocked at 800/ha consume 400-500 kg of Azolla daily gaining 800-1200 g in 5 months. In China this species is known to eat grass, leaves, small fishes, insects and other items in addition to aquatic vegetation or when such vegetation is in low supply. About half the plant food passing through the digestive system is undigested and large quantities of plant material must be eaten to sustain life. This consumption rate is the reason for its success at aquatic vegetation control (Greenfield, 1973). Grass carp overwinter without feeding.

Reproduction

A spawning migration to a large river takes place at about 15-17°C water temperature. The female swims in the centre of the river at the surface accompanied by 2-3 males, they roll and rub their bodies together and often jump out of the water. A male prods the female's body to stimulate egg release and leans closely to one side. Eggs are semi-buoyant and require a slow and steady current to keep them off the bottom (minimum water velocity of 0.23 m/second or more to support them and allow hatching; this is found in large rivers where the eggs hatch as they drift downstream; at 20°C and a not unusual velocity of 1.2 m/second, hatching requires 180 km of river). Temperatures should be above 20°C and preferably 21-25°C, or 26-30°C in another source. At these temperatures hatching takes about 40 hours. Flow rates should be 0.7-1.8 m/sec. Spawning occurs after heavy rain in rising rivers, when turbidity may reduce predatory attacks on the semi-pelagic eggs (Greenfield, 1973). This regime is also required for newly hatched fry and such conditions are rare outside their native habitat. In the Terek River of the Caspian Sea basin, the spawning migration begins in mid-April at water temperatures of 15-17°C and continues until August although numbers begin to decrease from the end of May. Spawning takes place after a sharp rise in water level and current speed. Eggs are first found in the drift in the second week of June and hatch 34-70 hours later depending on temperature. Some larvae reach rice fields and live there until autumn when the fields dry up, some being lost, others migrating. Other larvae are carried into the Caspian Sea where they are sensitive to the prevailing salinity at 1-1.5 days old (Abdusamadov, 1986). Up to 100,000 eggs are laid at one time (Greenfield, 1973) and in the Terek River fecundity reaches 1,230,700 eggs (Abdusamadov, 1986). Absolute fecundity may reach 2 million eggs. Eggs are up to 2.5 mm in diameter before fertilisation and are greyish-blue to bright orange. In water they swell to over 5.3 mm in 2 hours, becoming buoyant in flowing water.

Parasites and predators

Mokhayer (1976b) reports the cestode Bothriocephalus gowkongensis and the acanthocephalan Pomphorhynchus perforator. Red-sore disease is reported from fish pond grass carp in Iran by Razavilar et al. (1981). It is caused by a bacterium Aeromonas hydrophila and treatment was unsuccessful. Mokhayer (1989) records metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species, as well as shedder scales (sic), Echinochasmus perfoliatus. Jalali and Molnár (1990b) record the monogenean Dactylogyrus lamellatus from this species at fish farms in Iran. Viral haemorrhagic disease has been reported from grass carp in Iran (Iranian Fisheries Research and Training Organization Newsletter, 6:6, 1994; 9:6, 1995). Pond-cultured grass carp were found to be infected by the tapeworm Bothriocephalus, with 70-80 parasites causing intestinal obstruction and lowered haemoglobin, haematocrit and erythrocyte values (Esmaeli and Abbasi, 1996). Esmaeli and Peighan (1997) record an Aeromonas-like bacteria from grass carp in Khuzestan Province. Ebrahimzadeh Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. The crustacean parasite Lernaea elegans is reported from this species in the Mahabad Dam reservoir (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000). The intestinal helminth Bothriocephalus gowkongensis was recorded from this species on fish farms in West Azarbayjan Province (Azarvandi et al., 1999). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoan Ichthyophthirius multifilis, a copepod crustacean Lernaea sp., monogenean trematodes Dactylogyrus lamellatus, D. ctenopharyngodonis, and Gyrodactylus sp.. Jalali et al. (2002) and Jalali and Barzegar (2006) record Diplostomum spathaceum and Dactylogyrus lamellatus from this species in Lake Zarivar. Esmaeili et al. (2005) found a Flavobacterium columnaris-like bacterium on grass carp form Khuzestan fish ponds, suspected of either causing a 40% mortality or being a secondary factor in the fish kill. Pazooki et al. (2005) record Ergasilus peregrinus from this species in waterbodies of Zanjan Province. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus lamellatus from this species in the Mahabad River of the Lake Orumiyeh basin. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Trichodina sp., Dactylogyrus lamellatus, Lernaea cyprinacea and Diplostomum spathaceum. Barzegar et al. (2008) record the digenean eye parasites Diplostomum spathaceum and Tylodelphys clavata from this fish. Alishahi et al. (2009) examined moribund grass carp from 20 farms in Khuzestan for bacterial agents but found Aeromonas hydrophila, A. veroni and A. sohria in only 53 of 300 fish, secondary infections and not the cause of mortality. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Ergasilus sp., Ergasilus peregrinus, Lernaea sp. and Lernaea cyprinacea on this species. Maktabi et al. (2011) recorded the incidence of Listeria spp. on Khuzestan market fish from fish farms and the frequency was a matter of concern..

Any piscivore will take this species. Ashoori et al. (2012) found that grey herons (Ardea cinerea) in the Siahkeshim Protected Area of the Anzali Wetland ate this species.

Economic importance

This species has been introduced to Iran to control aquatic weeds in drainage and irrigation canals as an alternative to using polluting chemicals or mechanical removal. In some countries at is sought after by anglers. Grass carp may also help to control the snail-carried, human disease schistosomiasis, since the vegetation on which the snails live is severely reduced. They are also a food fish which relies on food sources not available to native fish (few fish consume whole plants). Grass carp consume vegetation at a rate of 100:1, i.e. for every 1 kg increase in grass carp biomass 100 kg of vegetation is consumed. Removal rates may exceed this figure since leaves are bitten off and branches clipped with not all of it being consumed. In the Dez Irrigation Project large amounts of this vegetation were removed daily from screens in test sections. Stocking in the Dez Irrigation Project in Khuzestan showed a removal rate approximately the same as mechanical control (Saadati, 1974; Behnke, 1975a). During a 5 month period the grass carp controlled 250 tons of aquatic vegetation per hectare. The fish preferred plant species which blocked the canals (Potamogeton spp. and Alisma gramineum) rather than those which grew close to the substrate (Chara and Cladophora) and did not interfere with water flow. Shireman and Smith (1983) give details on artificial propagation of this species.

Esmaeilzadeh et al. (2004) studied the nutrient composition and marinade qualities of this fish in Iran and compared them to those for safid mahi (Rutilus frisii) and found them to be preferable according to the organoleptic properties. The marinades could be stored for 6 months at 10ºC.

Fish farming of this species in Sistan was discontinued as its consumption of vegetation was reducing food for other species (www.netiran.com, downloaded 28 February 2005).

Holčík and Oláh (1992) report a catch of 315 kg in the Anzali Mordab in 1990. However Iran acounts for almost all the production of grass carp in the Near East and North Africa (4378 tonnes in 1994) (Food and Agriculture Organization, Fisheries Department, 1996). The aquaculture production in 1995 was 3942 tonnes (Bartley and Rana, 1998b). Grass carp sold for about U.S.$2.00/kg in 1995 (Rana and Bartley, 1998a). Marjan Iran Company was selling 1500-2000 g fish for U.S.$2.10/kg in August 2003 (http://groups.yahoo.com/groups/hilsa/message/25).

The inland waters of Turkmenistan had catches of 23 to 29.7 tonnes for the years 1971-1974 and a catch of 76 tonnes in 1970 when a ban on taking phytophagous fish was lifted.

Greenfield (1973) reviews the advantages and disadvantages of using this species as a weed control agent in the U.S.A. and Charyev (1984) in the Kara-kum Canal in Turkmenistan. Destruction of habitat for fishes and waterfowl, competition with native species and introduction of exotic diseases and parasites are all problems once this fish escapes into a main river suitable for reproduction. Their destruction of plants may interfere with waterfowl management, destroy breeding grounds for other species and facilitate the attacks of predators. Ideally triploids, produced by cold or warm shocks or by hydrostatic pressure on fertilised eggs, should be used initially as they cannot reproduce (Clugston and Shireman, 1987). However the chromosome number of each fish must be checked (by electronically measuring the volume of a red blood cell nucleus) as the process is not 100% effective. Grass carp are reproducing naturally in the Kara-kum Canal, vegetation is controlled, fish stocks have increased and some reduction of mosquitos has been obtained. However the ecosystem has been changed, spawning grounds of commercial species threatened, undesirable species have been introduced accidentally, and reduction in vegetation affects water quality. Grass carp are best used in restricted areas where improved flow and reduced mosquito populations are required but where there is no commercial fishery (Charyev, 1984).

The grass carp has a short gut and about half the plant material eaten is released to enrich the water and promote algal blooms. Oxygen levels and water clarity are reduced. The removal of plants can remove food sources for other fishes, shelter and spawning substrate. Additionally, as noted, the triploid treatment is not always effective and the species can become established.

Iran has had problems with disease outbreaks and poor survival of fingerlings which has led to production problems (Shehadeh, 1997).

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaculture, as food and in textbooks. There are numerous studies on this species as an experimental fish and in relation to aquaculture. Some Iranian studies include Alboughobish and Khaksari Mahabadi (2005) on the histology of the liver and pancreas; Pahn et al. (2005) used electrocardiograms to determine that the anaesthetic ketamine had no marked effect on heart activities; Morovvati et al. (2006) on seasonal changes of pronephros lymphoid tissue; Nahavandi et al. (2006) on the chemiluminescent response to determine the effect of various concentrations of diazinon, an organophosphate, on phagocytosis in order to measure immunity after exposure to this toxin; Pourgholam et al. (2006) and Sharifpour et al. (2006) on the toxicity and histopathological effects of diazinon; Sharifpour et al. (2006) on the sub-lethal effects  of diazinon on various organs; Pourgholam et al. (2006) on the toxicity of diazinon and the effects of sub-lethal concentrations on haematological and biochemical indices; Rezaei et al. (2007) on sensory evaluation and lipid quality of fish stored in ice - good to excellent until the fourth day and good to acceptable to the tenth day; Khajeh et al. (2008) on haematological parameters in cultured fish and found some to be lower than in Mesopotamichthys sharpeyi; Afkhami et al. (2011) on a comparison of chemical composition with Cyprinus carpio, there being significant differences in protein, lipid and moisture but not in ash; Niani and Khajeh-Ramhimi (2012) on the beneficial effect of gelatin coating as a preservative of refrigerated slices; Keramat Amirkolaie et al. (2010) on changes in gut morphology and growth in fish fed pellet and grass diets; Ghomi et al. (2011) on managing fertilization of fish ponds in order to manage teh elevsl of lernaeasis; Morovvati et al. (2011) on the histology of the excretory portion of the kidney; Makvandi et al. (2010) on successful culture in brackish water up to 3‰; Nekoubin and Soudagar (2012a) on the effect of formulate and plant diets on growth performance and survival rate of juveniles; Nekoubin and Soudagar (2012b) on growth and survival when a synbiotic is used in artificial diets; Pourgholam et al. (2013) on the haematological and biochemical changes in blood serum of fish vaccinated with Aeromonas hydrophila after exposure to diazinon, the agricultural pesticide, etc.

Conservation

No conservation is required for this exotic species. Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 10 million were produced in the Safid Rud Fish Farm in 1986. 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Feeding and growth studies on this species have also been carried out on this species in the Shaid Rajaee Hatchery (Ahmadi and Rezai, 1998). Experiments on induction of triploidy have been carried out in Iran using cold and heat shocks (M. Hassanzadehsaber, M. Pourkazemi, M. R. Nowruzfashkhami and A. Ghanaatparast (www.meeresschule.com/cgi-bin/abstracts/gastbuch.asp, downloaded 17 January2005).

Further work

Studies on the interactions of this species and native Iranian taxa should be carried out and introductions carefully controlled and monitored.

Sources

Shireman and Smith (1983) give a summary of the biology of this species. There is an extensive literature on herbivorous fishes, a recent book being Opuszynski and Shireman (1995), which has sections on grass carp. Gholipour (1996) has an account in Farsi.

Comparative material: BC65-381, 2, 95.4-98.8 mm standard length, Singapore, fish ponds (no other locality data).

Genus Cyprinion
Heckel, 1843

Scaphiodon Heckel, 1843 has been used for Cyprinion and Capoeta species in Southwest Asia.

Taki (1975) related members of this genus to a common ancestor with Onychostoma Günther, 1896, a Chinese and southeast Asian genus although Li et al. (2008) found this lineage to be unsupported on DNA evidence. Howes (1982) synonymises Semiplotus Bleeker, 1859, a genus found from Nepal to Viet Nam, and Scaphiodonichthys Vinciguerra, 1890, a genus from Indochina, with Cyprinion and refuted Taki's (1975) view using osteological characters, particularly of the jaws. Howes (1982) considers that Cyprinion cannot be defined on any uniquely derived characters. Krupp (1983) considers Howes' revision as unsatisfactory for the reasons that type specimens were not examined, relationships are based on jaw anatomy and other characters are largely excluded, variability of osteological characters within a species are largely unknown, and synapomorphies are not unequivocal. Bănărescu (1992b) and Banarescu and Herzig-Straschil (1995) regard Semiplotus as a distinct genus but probably related to Cyprinion. They comment that Semiplotus differs sharply from Cyprinion s.s. in the absence of barbels, a higher number of branched dorsal fin rays (20 or more), and in a lower number of branched anal fin rays (5 as in most related genera rather than the unusual 7 in Cyprinion). Scaphiodonichthys has 2 pairs of barbels (only 1 in Cyprinion), and 5 branched anal fin rays as well as differing from both Cyprinion and Semiplotus by having the lateral line closer to the ventral margin of the caudal peduncle and divergent rather than parallel striae on the scales. These latter 2 characters justify generic separation of Scaphiodonichthys. Bănărescu (1997) considers Scaphiodonichthys as valid and not a synonym of Cyprinion. Characters used by others to define Cyprinion such as expansion of the proximal part of the pelvic fin rays, interpelvic papillate flaps (Banister and Clarke, 1977) and a naked predorsal ridge (Mirza, 1969) do not occur in all species in this genus. If Semiplotus is included in Cyprinion then several osteological structures, particularly a synarthritic dentary joint, are uniquely derived or synapomorphic.

In the absence of a detailed revision, I have retained species within Cyprinion as the most familiar name in use in Southwest Asia for these fishes. Cyprinion s.s. is found from the Indus River basin west to the Arabian Peninsula and the Tigris-Euphrates basin but excluding northern drainages such as the Lake Orumiyeh, Caspian Sea and Hari River basins and excluding the westernmost edge of Southwest Asia such as the Jordan River basin and coastal drainages of Israel.

The genus Cyprinion is currently under revision by Florian Wicker at the Senckenberg Museum, Frankfurt and the status of the following species may undergo some changes.

Saadati (1977:45) refers to a new and undescribed Cyprinion species from Lar in southern Iran but the fish are Carasobarbus luteus.

A thorough study of the systematics of this genus in Iran depends to some degree on material from other areas which is not readily available, on large series of well-preserved adult specimens, and analyses which demonstrate consistency in characters used to define species. These conditions have not been achieved thus far in any studies undertaken and given the wide distribution and individual variation shown by Cyprinion species an adequate understanding of the species composition is not entirely possible.

This genus is characterised by a moderate sized, compressed body, a thick and blunt snout, an inferior mouth with a straight, crescentic or arched shape and a sharp horny edge to the lower jaw (which may fall off in preserved specimens), 1 pair of small barbels at the mouth corner, the last dorsal fin unbranched ray is thickened and bears weak to strong serrations (highly variable between individuals within a species and not a good character in species definitions), the dorsal fin is long (up to 16 branched rays) and the anal fin short (typically 7 branched rays), a ridge in front of the dorsal fin is formed internally from fused pterygiophores and lacks scales externally, pharyngeal teeth are in 3 rows and are compressed and spoon-shaped, scales large to moderate in size (lateral line counts (31-45), breast and belly scales may be absent (individually variable and not a good character), scale radii are restricted to the posterior field, peritoneum black, and gut very long and coiled (several times body length).

Cyprinion kais
Heckel, 1843


Dez River at Dez Wildlife Refuge, 24 April 2008, courtesy A. Mahjoor Azad
Dez River at Dez Wildlife Refuge, 24 April 2008, courtesy of A. Mahjoor Azad

Common names

butak-e dehan kuchek; بوتك (= botak); butak dahan kuchek, butok, لوتك (= lotak); smallmouth lotak; zanbour; زنبور دهان كوچك (= zanbour dahan kuchek).

[bunni saghir, bnaini; kais at Aleppo (= Haleb, Syria), hence the scientific name, all in Arabic; kais kingfish].

Systematics

Cyprinion Cypris Heckel, 1843 is a synonym, being a juvenile with keratinization of the lower jaw incomplete according to Howes (1982), although he did not examine the types. Krupp (1985c) and Banarescu and Herzig-Straschil (1995) agree with this synonymy. Berg (1949) placed C. kais (and C. cypris) in C. macrostomum, as the position of the dorsal fin in relation to the pelvic fins was variable in these fishes and not sufficient to warrant species status as Heckel (1843) stated in describing these species.

The type localities for Cyprinion Kais are "Aleppo" and "Mossul" and for Cyprinion Cypris the "Tigris bei Mossul" (Heckel, 1843b).

The syntypes of C. kais are in the Naturhistorisches Museum Wien comprising 3 fish in NMW 52801 (paralectotypes) and measuring 68.5-97.3 mm standard length, 2 fish in NMW 52802 measuring 120.6-164.3 mm standard length, and 2 fish in NMW 52803 (paralectotypes) measuring 153.4-154.2 mm standard length, the smaller of these being designated as the lectotype by F. Krupp in 1984. Eschmeyer et al. (1996) list possible syntypes in the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 2485 (2 fish, formerly NMW) and RMNH 2489 (1), and 1 syntype in the Senckenberg Museum Frankfurt (SMF 134, formerly NMW). The catalogue in Vienna lists 5 specimens.

A syntype of C. cypris is in the Senckenberg Museum Frankfurt (SMF 849, formerly NMW) (F. Krupp, pers. comm., 1985). Two syntypes, 63.5-106.2 mm standard length are under SMF 849, the larger one designated as a paralectotype (March 2007). Ten syntypes are in the Naturhistorisches Museum Wien (NMW 52804) measuring 51.2-115.1 mm standard length, the largest being designated as the lectotype (however Banarescu and Herzig-Straschil (1995) give 44.1-110.0 mm standard length for these 10 fish with one at 99.8 mm standard length as lectotype as selected by F. Krupp in 1984). Another specimen, 110.5 mm standard length, may also be a syntype (NMW 52800); and also NMW 59508, a dried specimen (Eschmeyer et al., 1996). The catalogue in Vienna lists 6 fish in alcohol and 1 fish stuffed.

Key characters

Mouth shape is distinctive. It is small and semicircular with a width about the size of the eye diameter and has large lateral lobes (= lower lips)(Kafuku, 1969). The cartilaginous sheath is thickened between the corners of the mouth and is rounded posteriorly with a distinct margin. The cartilage can form a tooth-like structure protruding anteriorly from the lower lip. The mouth in C. macrostomum is wider, arched and lacks the lateral lobes (see also illustrations in Kafuku (1969), Krupp (1985c) and Banarescu and Herzig-Straschil (1995)). These latter authors have the width of the mouth opening as only 13.5-22.0% of the head length (22.0-27.0% in C. macrostomum) for adult fish and the height of the arch or mouth opening (a line perpendicular from a line between the mouth corners to the tip of the lower jaw) 48-80% of the mouth width (29-47% in C. macrostomum), i.e. the mouth is narrower and more arched in C. kais. On this character, therefore, the two species can be distinguished as adults but there is potential for confusion in young fish. A single specimen identified as C. kais on the basis of mouth shape from the Dalaki River of Iran had values of 23.2% and 47.4% which are arguably C. macrostomum values. This specimen has a protruding tooth-like edge to the lower jaw in a u-shaped mouth with well-developed lips posterior to the "tooth".

The intestine is shorter and less complexly coiled in this species and the mean number of gill rakers is less in contrast to C. macrostomum (Kafuku, 1969). The back is higher and more curved, the eyes are larger and the anal fin is more posterior, in addition to the mouth shape (Heckel, 1843b). The dorsal fin origin arises over that of the pelvic fins (Heckel, 1847a). The edge of the dorsal fin is more notched in C. kais than in C. macrostomum (the length of the fourth branched ray is 48-62% of the length of the first ray as opposed to 55-79% in C. macrostomum, with extreme values overlapping, according to Banarescu and Herzig-Straschil (1995)).

The form of the pharyngeal teeth is different from C. macrostomum (see Krupp (1985c) for illustrations where kais has hooked tips and macrostomum does not), there are fewer gill rakers (8-12 on the lower arch in kais, 12-16 in macrostomum), on average there are fewer dorsal fin rays, the last unbranched dorsal fin ray is longer, and interorbital width is smaller. However sample sizes in some studies are small (in Kafuku (1969) only 5 fish of each species were examined), morphometric characters are notoriously size-dependent, gill raker counts are also size dependent, and even pharyngeal tooth form varies with age (small macrostomum have hooked tips). C. kais may well be a good species but a wide-ranging comparison of adults and young and of localities is needed and material from Iran is scarce or equivocal. Further discussion is under C. macrostomum.

Morphology

Dorsal fin with 4 unbranched and 12-16 branched rays, anal fin with 3 unbranched and 7 branched rays. The dorsal fin has the last unbranched developed as a spine with strong teeth except at the extreme tip which is thin and flexible. Pectoral fin with 14-18 branched rays, and pelvic fin with 8-9 branched rays. Lateral line with 36-43 scales. The belly is scaled. There is a well-developed pelvic axillary scale. Scales have a subcentral anterior focus, fine circuli, few posterior radii and no or very few anterior radii. Total gill rakers 10-15, short and reaching the raker below when appressed. Rakers are absent on the anterior arch where there are only tubercles. Pharyngeal teeth 2,3,4-4,3,2, with variants 2,3,5-5,3,2 and 2,3,5-4,3,2, spoon-shaped with a small hook at the tip. Chromosome number is 2n=50 (Nasri et al., 2010).

Meristics for Iranian material: dorsal fin branched rays 12(2), 13(2), 14(1) or 15(1); branched anal fin rays 7 (6); branched pectoral fin rays 14(1), 15(2), 16(2) or 17(1); branched pelvic rays 8(4) or 9(2); lateral line scales 38(5) or 39(1); and total gill rakers 12(2), 13(3) or 14(1).

Sexual dimorphism

Tuberculation in a 103.5 mm standard length specimen consisted of ca. 20 tubercles restricted to the area over the lachrymal bone. A specimen 147.5 mm standard length had small to minute tubercles in front of the eye, under the eye, on the mid-preoperculum and on the mid-operculum. Curiously the individual small tubercles on the operculum were connected by thin lines of horny tissue.

Colour

Overall colour is silvery to yellowish-white with the back grey-brown and the lower surfaces a lemon yellow. The lower jaw margin is a glossy yellow. The fish shown above may represent a spawning colouration, not seen in all specimens. The pelvic fins are a bright orange-red, the pectorals paler. Some fish have a less strong colour in the pelvic than in the anal fin. The anal fin is yellow, to orange or greenish, distally black and anteriorly most orange. The caudal fin has light orange to greenish tints. The dorsal fin is black with a yellow-tinged base becoming anteriorly reddish. In preserved fish, there is some concentration of pigment above and below each lateral line pore, scales on the back and upper flank are outlined with pigment, and there is some concentration of pigment into a few to moderate number of diffuse spots on the uppermost flank and back midline. The leading edge of the dorsal fin is very dark (but may be light), dorsal fin membranes are dark, anal fin membranes also dark but to a lesser extent, and the caudal, pectoral and pelvic fins have pigment lining the rays. Peritoneum black.

Size

Attains 21.5 cm total length, or to 25.0 cm total length in Iraq (Al-Rudainy, 2008).

Distribution

This species is found in the Tigris-Euphrates and Quwaiq basins. Abdoli (2000) maps the Jarrahi, Karun, middle to lower Dez, and Karkheh up to the Simarreh rivers of the Tigris River basin. It is present in in the Niloufar spring, Kermanshah (Eagderi and Nasri, 2012). It is also found in the Gulf basin, although rare, and specimens from the sugar cane fields of Khuzestan were seen in 2000 (personal observations, B. W. Coad).

Zoogeography

Zoogeographical comments are under the genus above.

Habitat

This species is recorded from a variety of habitats as listed above and is also known to inhabit canals but nothing is known of its environmental requirements.

Age and growth

Unknown.

Food

Gut contents are filamentous algae in the one specimen examined. Diet may be similar to Cyprinion macrostomus. Al-Rudainy (2008) gives aquatic insects and detritus for Iraq. Curiously, the mouth structure resembles that of the unrelated cutlips minnow, Exoglossum maxillingua (Le Sueur, 1817), from North America. This species feeds on insect larvae, with some molluscs and worms. Food is scraped from the bottom or poked out of crevices using the shovel-like lower jaw. Sand is also taken in and spat out, presumably after food items are extracted. The cutlips also picks out the eyes of other fishes in confined areas (Coad et al., 1995).

Reproduction

Generally unknown. Ünlü (2006) gives age at first maturity as 2 years in the Turkish Tigris River with spawning over sand, stones and gravel in May-June.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

This minnow appears to be rare, or at least is rarely collected, in Iran. Cyprinion macrostomum is much more common and is taken in most seine hauls in streams and rivers. The distribution and population numbers are unknown. Endangered in Turkey (Fricke et al., 2007).

Further work

The biology of this species needs to be investigated and the use of the peculiar jaw structure ascertained. Its great rarity, at least in Iran, leads to the suspicion that it may be a developmental anomaly of Cyprinion macrostomum - the few specimens at hand don't permit a detailed study of characters other than the strikingly different jaw (see comments under C. macrostomum and also above). Development of pharyngeal teeth, gill raker numbers, complexity of gut coils and morphometric characters are all size dependent and show individual and populational variations not analysable here.

Sources

Type material: See above, Cyprinion kais (NMW 52801, 52802 and 52803) and C. cypris (NMW 52804).

Iranian material: CMNFI 1993-0141, 1, 66.3 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); CMNFI 2008-0169, 5, 80.4-98.2 mm standard length, Khuzestan, irrigation ditch in sugar cane fields (31º58'42"N, 48º31'07"E); ZSM 25715, 2, 34.1-65.3 mm standard length, Khuzestan, Dez River at Harmaleh (31º57'N, 48º34'E).

Comparative material: BM(NH) 1920.3.3:50, 1, 83.6 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1920.3.3:94-115, 40, 65.3-92.4 mm standard length, Iraq, Basrah (30º30'N, 47º47'E); BM(NH) 1931.12.21:3, 1, 129.8 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:115-120, 5, 90.6-147.9 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1105, 1, 115.6 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1106, 1, 101.4 mm standard length, Iraq, Fao (29º58'N, 48º29'E); BM(NH) 1974.2.22:1214-1255 (in part), Iraq, Khalis (33º49'N, 44º32'E); BM(NH) 1984.4.18:30, 63.4 mm standard length, Iraq, Kut Hiwa (no other locality data); FMNH 51229, 1, 103.5 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); FMNH 51230, 6, 42.9-60.5 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); FMNH 51231, 2, 64.0-64.8 mm standard length, Iraq, Diyala River, 12 miles east of Baghdad (no other locality data); uncatalogued, 5, 49.1-66.7 mm standard length, Iraq, Shatt al Arab (no other locality data); uncatalogued, 1, 107.2 mm standard length, Turkey, Euphrates River 20 km west of Erzurum (ca. 41º03'N, ca. 39º55'E).

Cyprinion macrostomum
Heckel, 1843

Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi
Gamasiab River, Karkheh River basin, January 2010, courtesy of K. Abbasi

Common names

بوتك (= botak); butok; لوتك (= lotak); largemouth lotak; butak-e dehan (or dahan) buzorg in Khuzestan; galuk (Mokhayer (1981c); kapour; zanbour (= bee) in Khuzestan and Boyer Ahmadi-ye Sardsir va Kohkiluyeh provinces; زنبور دهان بزرگ (= zanbour dahan bozorg); ?tumbuek (= hunting horn, possible name from Heckel (1843b)).

[hmarriya sefra or himriya sefra, surrah masih, bunni kaper, dunbuk kabir al-fam, benayne; kais at Aleppo (= Haleb, Syria) but see above species (Heckel, 1843b); dombok or dumbek at Mosul (= solid or compact flesh, a good source of food, according to Heckel (1843b)); all preceding in Arabic; large-mouthed barb, Tigris kingfish].

Systematics

Originally spelt macrostomus but correctly macrostomum (Berg, 1949). Cyprinion neglectus Heckel, 1847 from the "Tigris bei Mossul" is a synonym (Krupp, 1985c; Banarescu and Herzig-Straschil, 1995). Howes (1982) considered that Cyprinion tenuiradius (q.v.) was only a "variant" of this species but did not examine any material. Berg (1949) places C. kais (q.v.) in the synonymy of this species along with C. cypris (see C. kais).

The type locality of Cyprinion macrostomus is given by Heckel (1843b) as "Aleppo" and "Mossul". Krupp (1985c) lists 5 syntypes from Aleppo, 81-133 mm standard length in the Naturhistorisches Museum Wien (NMW 52805), the largest being selected as the lectotype (hence Aleppo is the type locality as designated by the publication of Banarescu and Herzig-Straschil (1995)). One syntype from Aleppo, 83 mm standard length, is in the Senckenberg Museum Frankfurt (SMF 70, formerly NMW; Eschmeyer et al. (1996) give SMF 870) and 4 syntypes from Mosul, 58-124 mm standard length are in the Naturhistorisches Museum Wien (NMW 52806). My measurements are 82.1-135.0 mm standard length for NMW 52805 and 59.1-126.2 mm standard length for NMW 52806. Another syntype is a dried specimen NMW 52503, and the Rijksmuseum van Natuurlijke Historie, Leiden has 1 syntype under RMNH 2487, formerly NMW) and 1 syntype under RMNH 2488, formerly NMW). The catalogue in Vienna lists 4 specimens.

Seven syntypes of Cyprinion neglectus from Mosul measure 54-131 mm standard length (NMW 52807), the largest being selected as the lectotype (Krupp, 1985c). My measurements are 53.3-131.9 mm standard length (Banarescu and Herzig-Straschil (1995) have 53.1-128.2 mm standard length). All material was collected by Th. Kotschy in 1842 for Aleppo and 1843 for Mosul. The catalogue in Vienna lists only 2 specimens under this name.

?Check lengths against data sheets

Daştan et al. (2012) examined the genetic diversity of this species in Anatolia and compared it with C. kais.

Key characters

Distinguished from C. kais by mouth and dorsal fin ray characters as described under that species, by having more gill rakers and a longer and more coiled intestine (Kafuku, 1969). The dorsal fin origin is in front of that of the pelvic fins (Heckel, 1847a). See discussion under C. tenuiradius for distinction from that taxon.

Morphology

Dorsal fin with 4 unbranched and 12-17 branched rays (usually 14-15 according to Banarescu and Herzig-Straschil (1995) but 77% of fish in Iran are 13-14, see below). The last dorsal fin unbranched ray is strong and serrated to the tip. The anal fin has 3 unbranched and 6-7, usually 7, branched rays. In Iranian specimens, 96.1% of 127 fish have 7 rays, the remainder 6 rays. Pectoral fin branched rays are 12-17 and pelvic fin branched rays 7-9, usually 8. Lateral line scales 33-45 (usually 41-44 according to Banarescu and Herzig-Straschil, 1995) but a broader range in Iran, see below). The breast is covered with scales. The pelvic axillary scale is very elongate. Scales are squarish, being deeper than long, often with parallel dorsal and ventral margins (or rounded margins). The anterior margin has a marked central protuberance and the posterior margin is rounded. Radii are numerous on the posterior field and circuli are fine and numerous. The posterior field circuli break into "bubbles". The focus is subcentral anterior. Gill rakers 16-17, on the lower arm 12-16, in the literature but a much wider range in total rakers in Iran (see below). Rakers are short and only touch the raker below or a little further when appressed. Pharyngeal teeth 2,3,5-5,3,2, 2,3,4-4,3,2, and variations on 4 or 5 main row teeth. Teeth are spatulate with broad, flattened crowns. The tips of teeth are slightly hooked in small fish. The most anterior tooth in the main row may be very small or absent (or incompletely ossified and hard to distinguish). The gut is very elongate with complex coils. In small fish, the upper lip is not covered with a fold of the snout as in large fish. Also the gut is not as coiled in young fish as in adults. Chromosome number is 2n=48 (Ünlü et al., 1997; Gaffaroğlu and Yüksel, 2004).

Meristics for Iranian fish from the Tigris River basin: branched dorsal fin rays 12(4), 13(43), 14(52), 15(26) or 16(3)(mean = 13.9, S.D. = 0.861); branched pectoral fin rays 14(3), 15(44), 16(57) or 17(25)(mean = 15.8, S.D. = 0.771); branched pelvic fin rays 7(7), 8(121) or 9(1)( mean = 8.0, S.D. = 0.246); lateral line scales 33(3), 34(1), 35(12), 36(11), 37(3), 38(11), 39(29), 40(31), 41(25), 42(2) or 45(1)(mean = 38.8, S.D. = 2.211); total gill rakers 13(3), 14(8), 15(15), 16(23), 17(15), 18(24), 19(17), 20(14) or 21(6) (mean = 17.3, S.D. = 2.022); pharyngeal teeth 2,3,5-5,3,2(17), 2,3,4-5,3,2(8), 2,3,5-4,3,2(3), or 2,3,4-4,3,2(2); and total vertebrae ?.

The mouth is usually transverse or slightly arched and usually has a horny covering. Small fish have a crescentic mouth. A wide range of mouth arching is seen in fish of varying sizes and even in fish of the same size and locality of capture. Banarescu and Herzig-Straschil (1995) note that the syntypes of Cyprinion neglectus have a mouth arch which is more curved and not as wide, somewhat intermediate between C. macrostomum and C. kais, being closer to the former. This variation is attributed to the material possibly being from some tributary of the Tigris River, or from isolated ponds, where introgression with C. kais took place. It may well be that variation in mouth shape is more marked than limited sample sizes would indicate. Certainly in smaller fish, e.g. in 20 specimens of C. macrostomum (38.5-54.0 mm SL) examined by me from Iran, values for mouth width and depth as measured in Banarescu and Herzig-Straschil (1995) are not as clear cut and there is a variable developmental gradient in mouth shape. Mouth "height" as a % of width was 29.2-53.8 and width as % of head length was 22.1-36.6. Banarescu and Herzig-Straschil (1995) give "height" as 19-31% of width and width as 26-44% head length for macrostomum and 48-80% and 13.5-22.0% respectively for kais. Large macrostomum and kais (>100 mm SL) can be distinguished on mouth shape but not smaller specimens which bridge the gap between the two species. The possibility that kais is a developmental anomaly of macrostomum, retaining juvenile features, should be investigated.

Sexual dimorphism

Mature males have large tubercles on the snout in a broad band below the nostril level, extending back under the eye and breaking up into a few tubercles on the operculum. There is a large tubercle between the nostril and the eye. Fine tubercles are scattered over the top of the head. Three tubercles are found in rows on the first branched pectoral fin ray and very strong tubercles line each anal fin branched ray in single file. The anterior pelvic fin rays have the occasional 1-2 tubercles or a row of tubercles. Dorsal and caudal fin rays have fine tubercles, much smaller than those on the anal fin. Mid and posterior flank scales have 1-3 small tubercles, variably arranged on the exposed scale.

Colour

The back is bluish-grey to bluish-black or brown, flanks silvery or silvery-yellow and the belly whitish with silvery tints. The upper head is light brown. Scales are outlined with dark pigment and the anterior exposed scale base is darkened. The cleithrum area is pink or orange in some fish with pink or orange spots on up to 5 rows of flank scales but mostly along the anterior lateral line. Fish from a saline stream in Khuzestan had a pale-pink cleithrum and lateral line spots. There is a reddish-yellow spot at the base of the pectoral and pelvic fins. The pectoral, pelvic, anal and caudal fins are yellowish to pinkish or orange proximally and blackish distally. The dorsal fin has a narrow, yellow stripe at the base and the rest of the fin is black. The cartilaginous lower jaw is reddish-yellow to orange. The eye is slightly yellow. Small live fish are silvery overall with a white belly and olive back, the pectoral and pelvic fins slightly orange-yellow and other fins greyish although all fins may be hyaline. The peritoneum is black.

Small preserved fish have an indistinct blotch at the caudal fin base and a similar blotch on the back at the base of the spine in the dorsal fin. In very small fish these blotches are more distinct and there are 4-7 irregular blotches on the mid-flank above the lateral line and 3 blotches at the dorsal fin base. Development of blotches is individually variable, some fish being almost immaculate while in others the blotches extend vertically as bars as far as the back.

Size

Reaches 19.3 cm standard length (Krupp, 1985c).

Distribution

Found in the Orontes, (= Asi), Quwayq and Tigris-Euphrates basins. In Iran, it is found in the Tigris River basin including the Hawr Al Azim, Khersan, Jarrahi, Marun and Gamasiab rivers (Berg, 1949; Abdoli, 2000; K. Abbasi, see photograph above) and the northern Gulf basin in the Shapur, Dalaki and Helleh rivers (Gh. Izadpanahi, pers. comm., 1995; Sedaghat and Hoseini, 2012c), the Zohreh River and possibly Lake Famur - some may be C. tenuiradius. Vossoughi (1998) reports this species from the western Hamun-e Jaz Murian basin based on a fishes with 13-15 branched dorsal fin rays, much higher than for C. watsoni, the taxon to be expected in this area.

Zoogeography

Zoogeographical comments are under the genus above.

Habitat

Known from a variety of habitats such as rivers, streams, reservoirs and ponds, as well as canals and gravel pits. Al-Habbib and Al-Habbib (1979) have demonstrated experimentally for a sample from "Nawaran Spring" north of Mosul, Iraq that this species can survive temperatures up to about 37°C. Akpinar and Aksoylar (1989) and Akpinar (1999) report this species from the Kangal Thermal Spring, Sivas, Turkey at a constant temperature of 35°C. This is the commonest species in catches in southwestern Iran, followed by Garra rufa. In areas under human influence in Lorestan, such as the lower reaches of rivers and near cities, it exceeds 80% in numbers in catches.

Age and growth

Maximum age reported for a population in the "Al-Nibaey" Lakes near Baghdad is 7+ years. Growth is slow and there is no difference in growth between males and females, although the habitat is not considered ideal for these fishes. Females tend to be slightly heavier than males of the same length especially in older fish. The length-weight relationship was W = 0.027 L2.67 (r = 0.78) for both sexes, W = 0.028 L2.65 (r = 0.90) for males and W = 0.020 L2.78 (r = 0.93) for females. Maturity is attained at 10.0-11.1 cm, corresponding to age group 2 (Allouse et al., 1989). The length-weight equation for commercially caught fish in the Tigris River was log W = 2.884 log L-4.623, condition factor was 1.15-1.47 (mean 1.28) and fish were immature up to age 2+ (Al-Nasiri, 1991). Haematology of this species from Sarao Subhana Agha near Sulaymaniyah was examined by Al-Mehdi and Khan (1984).

Sedaghat and Hoseini (2012c) found positive allometric growth for fish in the Dalaki River, Bushehr with the b value not significantly different between sexes and W = 0.06L3.10 for the population.

Food

Major food items in the Baghdad study are of plant origin with occasionally some chironomid larvae, copepods and cladocerans. Khan (1988) found for fish from near Sulaimaniyah, Iraq that diatoms and decayed organic matter are the main foods, with some green algae. Zooplankton are thought to be accidental food items. Guts contain mud and sand, evidence of a bottom feeding habit. Feeding increases at the start of the breeding season. The horny lower jaw covering is used to scrape algal food off hard bottom objects.

Reproduction

Near Baghdad, most fish are mature by April, the gonads occupying about one-third of the body cavity. Ovaries are orange to yellowish and testes milky white. Spawning occurs principally in May and June, with some in early July, but by July most fish are spent. Al-Rudainy (2008) gives a spawning season of May and June in Iraq on gravel beds in shallow water with fast current. Maturity is attained there at 2-3 years, 15 cm length and 50 g weight.

Iranian material shows minute but developing eggs in a 71.3 mm standard length fish caught on 31 January and specimens caught on 5 July have eggs 1.4 mm in diameter. The 31 January fish has tubercles on the snout and anal rays so tubercles develop quite early and in small fish. A fish caught on 20 September also shows tubercles around the snout. Small fish caught in January about 20 mm SL are presumably the young from the previous season and so show slow growth or are evidence of a prolonged or late spawning season.

Parasites and predators

Gussev et al. (1993a) describe a new species of monogenean from C. macrostomum in the Karun River, Dactylogyrus cyprinioni, and Jalali (1992) a new species of monogenean, Dogielius molnari, in the Dez River, both in Khuzestan. Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus pallicirrus, from fish taken in the Dez River near Ahvaz.

Economic importance

Al-Mehdi and Khan (1984) report this species to be important in riverine and culture fisheries in northern Iraq. Ündar et al. (1990) identify this species and Garra rufa as the "doctor fish" of the Kangal hot spring in Turkey (Timur et al., 1983; Warwick and Warwick, 1989; Kürkçüoğlu and Öz, 1989; and various newspaper and television reports). High water temperatures reduce the amount of plankton available as fish food and the fish nibble away infected skin of humans who bathe in these waters. The fish is known as "striker" (and Garra rufa as "licker") from its behaviour in the spa pools. The healing properties are linked to the high level of selenium (1.3 p.p.m.) in the water, selenium being beneficial in some skin diseases, and possibly to UV light. The fish facilitate the action of the selenium and UV light by softening and clearing away psoriatic plaque and scale, exposing the lesions to the water and sunlight. However, some lesions are made worse and the fish can cause some new ones.

Conservation

This species is widely distributed in southern areas, particularly Khuzestan, and does not appear to be under threat other than that suffered by all species by pollution and water abstraction. Endangered in Turkey (Fricke et al., 2007).

Further work

See comments above on the need for further work to distinguish this species from C. kais, especially when young and below for distinction from C. tenuiradius.

Sources

Type material: See above, Cyprinion macrostomum (NMW 52805, 52806), C. neglectus (NMW 52807).

Iranian material: Tigris basin: and presumably macrostomum CMNFI 1979-0268, 13, 92.2-122.4 mm standard length, Lorestan, Dez or Karkheh drainage between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0269, 4, 104.7-110.6 mm standard length, Lorestan, Dez or Karkheh drainage between Nowqan and Khorramabad (no other locality data); CMNFI 1979-0270, 10, 85.5-122.4 mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0271, 3, 100.7-144.8 mm standard length, Lorestan, Kashkan River drainage (33º39'N, 48º32'30"E); CMNFI 1979-0273, 9, ? mm standard length, Lorestan, Kashkan River drainage (33º26'N, 48º19'E); CMNFI 1979-0274, 14, ? mm standard length, Lorestan, Kashkan River drainage (33º27'N, 48º11'E); CMNFI 1979-0275, 2, 142.4-165.0 mm standard length, Lorestan, Kashkan River drainage (33º25'N, 47º58'E); CMNFI 1979-0278, 4, 93.5-114.1 mm standard length, Lorestan, Kashkan River drainage (33º34'N, 48º01'E); CMNFI 1979-0279, 9, 100.3-149.4 mm standard length, Lorestan, Khorramabad River (33º37'N, 48º18'E); CMNFI 1979-0283, 5, 93.0-144.0 mm standard length, Kermanshahan, Qareh Su drainage (34º21'N, 47º07'E); CMNFI 1979-0287, 1, 112.6 mm standard length, Kermanshahan, Chashmeh Javari 2 km from Ravansar (ca. 34º42'N, ca. 46º40'E); CMNFI 1979-0288, 1, 94.3 mm standard length, Ilam and Poshtkuh, Gangir River at Juy Zar (33º50'N, 46º18'E); CMNFI 1979-0289, 4, ? mm standard length, Kermanshahan, Diyala River drainage (34º28'N, 45º52'E); CMNFI 1979-0290, 11, 49.3-133.0 mm standard length, Kermanshahan, Diyala River drainage at Qasr-e Shirin (34º31'N, 45º35'E); CMNFI 1979-0291, 15, ? mm standard length, Kermanshahan, Diyala River drainage (34º24'N, 45º37'E); CMNFI 1979-0350, 18, ? mm standard length, Khuzestan, Marun River near Marun (30º39'30"N, 50º02'E); CMNFI 1979-0355, 1, ? mm standard length, Khuzestan, stream tributary to Karun River at Salmaneh (30º35'N, 48º22'E); CMNFI 1979-0356, 1, ? mm standard length, Khuzestan, stream at Hoveyzeh (31º27'N, 48º04'E); CMNFI 1979-0360, 2, ? mm standard length, Khuzestan, canal branch of Karkheh River (31º40'N, 48º35'E); CMNFI 1979-0361, 3, ? mm standard length, Khuzestan, jube in Karkheh River drainage (31º42'N, 48º33'E); CMNFI 1979-0363, 1, ? mm standard length, Khuzestan, Karkheh River (31º52'N, 48º20'E); CMNFI 1979-0364, 2, ? mm standard length, Khuzestan, river at Abdolkhan (31º52'30"N< 48º20'30"E); CMNFI 1979-0365, 24, ? mm standard length, Khuzestan, stream in Doveyrich River drainage (32º25'N, 47º36'30'E); CMNFI 1979-0366, 16, ? mm standard length, Khuzestan, stream west of Dehloran (32º45'30"N, 47º05'30"E); ID? CMNFI 1979-0367, 2, ? mm standard length, Khuzestan, Meymeh River 11 km north of Dehloran (32º44'30"N, 47º09'30"E) ID? CMNFI 1979-0368, 12, ? mm standard length, Khuzestan, Karkheh River (32º24'30"N, 48º09'E); CMNFI 1979-0371, 1, ? mm standard length, Khuzestan, stream in Karkheh River drainage (32º05'N, 48º19'E); CMNFI 1979-0373, 12, ? mm standard length, Khuzestan, Bala River north of Andimeshk (32º35'N, 48º17'E); CMNFI 1979-0374, 46, ? mm standard length, Khuzestan, stream tributary to Bala River (32º40'N, 48º15'E); CMNFI 1979-0376, 9, ? mm standard length, Khuzestan, river tributary to Karkheh River (32º48'30"N, 48º04'30"E); CMNFI 1979-0378, 10, ? mm standard length, Khuzestan, stream tributary to Karkheh River (ca. 32º48'N, ca. 48º04'E); CMNFI 1979-0379, 11, ? mm standard length, Khuzestan, Dez River (32º12'N, 48º27'E); CMNFI 1979-0380, 5, ? mm standard length, Khuzestan, stream tributary to Dez River (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0381, 28, ? mm standard length, Khuzestan, stream 40 km west of Shushtar (ca. 32º10'N, ca. 48º35'E); CMNFI 1979-0382, 67, ? mm standard length, Khuzestan, Karun River at Shushtar (32º03'N, 48º51'E); CMNFI 1979-0383, 1, ? mm standard length, Khuzestan, stream in Ab-e Shur drainage (31º59'30"N, 49º06'E); CMNFI 1979-0384, 7, 86.3-152.2 mm standard length, Khuzestan, Ab-e Shur drainage (32º00'N, 49º07'E); CMNFI 1979-0386, 4, ? mm standard length, Khuzestan, stream 21 km from Haft Gel (ca. 31º34'N, ca. 49º23'E); CMNFI 1979-0387, 6, ? mm standard length, Khuzestan, stream 12 km from Haft Gel, Jarrahi River drainage (31º25'N, 49º38'E); CMNFI 1979-0388, 2, ? mm standard length, Khuzestan, Zard River (31º19'N, 49º44'E); CMNFI 1979-0390B, 23, 36.2-156.2 mm standard length, Khuzestan, stream 3km south of Bagh-e Malek (31º29'N, 49º54'30"E); CMNFI 1979-0391, 1, 154.5 mm standard length, Khuzestan, stream in Marun River drainage (31º28'N, 49º51'E); CMNFI 1979-0392, 5, ? mm standard length, Khuzestan, Zard River (ca. 31º32'N, ca. 49º48'E); CMNFI 1979-0393, 2, 96.9-116.6 mm standard length, Khuzestan, Jarrahi River drainage (31º18'N, 49º37'E); CMNFI 1979-0394, 1, 130.2 mm standard length, Khuzestan, stream in Marun River drainage (31º01'N, 49º45'E); CMNFI 1979-0395, 4, ? mm standard length, Khuzestan, stream in Marun River drainage (ca. 30º57'N, ca. 49º51'E); CMNFI 1979-0396, 1, ? mm standard length, Khuzestan, Kheyrabad River (30º32'N, 50º23'30"E); ID? CMNFI 1979-0398, 23, ? mm standard length, Boyer Ahmadi-ye Sardsir va Kohkiluyeh, stream in Zohreh River drainage (30º24'30"N, 50º37'30"E); ID? CMNFI 1979-0399, 7, ? mm standard length, Fars, stream in Zohreh River drainage (30º19'30"N, 51º15'E); CMNFI 1991-0153, 1, 171.3 mm standard length, Khuzestan, Zohreh River (no other locality data); CMNFI 1991-0154, 1, 109.9 mm standard length, Khuzestan, Hawr al-Azim (ca. 31º45'N, ca. 47º55'E); CMNFI 1993-0128, 1, 110.7 mm standard length, Kermanshahan, Sarab-e Sabz 'Ali Khan (34º25'N, 46º32'E); CMNFI 1993-0149, 1, 121.7 mm standard length, Khuzestan, Karun River (no other locality data); CMNFI 2007-0111, 6, 24.7-173.8 mm standard length, Kermanshahan, Alvand River near Sar-e Pol-e Zahab (ca. 34º36'N, ca. 45º56'E); CMNFI 2007-0112, 6, 46.5-118.8 mm standard length, Kermanshahan, Kerend River basin near Shahabad-e Gharb (ca. 34º06'N, ca. 46º30'E; CMNFI 2007-0113, 1, 122.1 mm standard length, Kermanshahan, Razavar River, Qareh Su tributary (ca. 34º25'N, ca. 47º01'E); CMNFI 2007-0115, 6, 59.7-154.8 mm standard length, Kermanshahan, Qareh Su basin (ca. 34º34'N, ca. 46º47'E); CMNFI 2007-0116, 12, ?-93.0 mm standard length, Kermanshahan, Gav Masiab basin west of Sahneh (ca. 34º28'N, ca. 47º36'E); CMNFI 2007-0117, 1, ? mm standard length, Kermanshahan, Gav Masiab basin near Sahneh (ca. 34º24'N, ca. 47º40'E); BM(NH) 1980.8.28:1, 1, 90.3 mm standard length, Khuzestan, Dezful (32º23'N, 48º24'E); BWC95-20, 14, ? mm standard length, Khuzestan, Rud Zard at Rud Zard (31º22'N, 49º43'E); Gulf fish:- ? tenuiradius CMNFI 1979-0020, 56, ?, mm standard length, Fars, Mand River outside Kavar (29º11'N, 52º41'E); CMNFI 1979-0054, 14, 37.4-64.1 mm standard length, Fars, Shur River tributary (ca. 28º58-29º03'N, ca, 52º34-35'E); CMNFI 1979-0075, 123, 21.3-142.4 mm standard length, Fars, Mand River at Pol-e Kavar (29º11'N, 52º41'E); CMNFI 1979-0109, 5, 63.2-100.2 mm standard length, Fars, Mand River at Shahr-e Khafr (28º56'N, 53º14'E); CMNFI 1979-0128, 7, 19.2-103.8 mm standard length, Shur River (28º51'N, 52º31'E); CMNFI 1979-0131, 19, 16.4-41.7 mm standard length, Fars, Ab-Arak River (28º38'N, 52º49'E); CMNFI 1979-0132, 72?, 15.2-100.1 mm standard length, Fars, Ab-Arak River (28º35'N, 52º58'E); CMNFI 1979-0133, 50, 45.6-95.5 mm standard length, qanat stream near Qir (28º27'30"N, 53º03'E); CMNFI 1979-0135, 18, 21.8-49.2 mm standard length, Mand River tributary (28º08'N, 53º10'E); CMNFI 1979-0157, 4, 23.6-85.4 mm standard length, Fars, qanat stream at Hadiabad (28º52'N, 54º13'E); macrostomum? CMNFI 1979-0193, 1, 36.3 mm standard length, Fars, river 8 km from Darab (28º45'N, 54º27'30"E); macrostomum? CMNFI 1979-0195, 1, ? mm standard length, Fars, jube west of Darab (ca. 28º54'N, ca. 53º53'30"E); CMNFI 1979-0196, 1, 59.9 mm standard length, Fasrs, qanat and pool at Khanehnehrin (28º50'N, 53º31'30"E); not on data sheet check jar? CMNFI 1979-0197, 1, 51.3 mm standard length, Fars, spring nd stream 33 km from Fasa (28º45'N, 53º25'E); CMNFI 1979-0198, 23, 22.3-57.7 mm standard length, Fars, stream at Tadovan (28º47'N, 53º24'30"E); CMNFI 1979-0200, 8, 29.0-46.1 mm standard length, Fars, Mand River tributary (28º36'N, 53º36'30"E); CMNFI 1979-0202, 12, ? mm standard length, Fars, Mand River (29º01'N, 53º00'E); CMNFI 1979-0241, 18, 43.8-72.6 mm standard length, Fars, Shapur River at Shapur (29º47'N, 51º35'E); CMNFI 1979-0347, 2, 105.2-106.7 mm standard length, Fasr, Pol-e Berengie (29º27'30"N, 52º32'E); CMNFI 1979-0348, 4, 52.9-79.1 mm standard length, Fars, stream at Somduldul (ca. 29º28'N, ca. 52º32'E); CMNFI 1979-0404, 25, 20.2-127.9 mm standard length, Bushehr, stream 33 km south of Kaki (28º08'N, 51º47'E); CMNFI 1979-0405, 4, 33.5-36.7 mm standard length, Hormozgan, stream about 13 km north of Rostaq (28º29"N, 54º59'E); ID? CMNFI 1979-0497, 1, 85.6 mm standard length, Fars, Mand River at Band-e Bahman (29º11'N, 52º40'E); CMNFI 1979-0501, 17, 18.7-91.0 mm standard length, Fars, Mand River at Kavar (29º11'N, 52º41'E); CMNFI 1979-0504, 6, ?-93.0 mm standard length, Fars, stream at Pol-e Gaz in Lake Maharlu basin (no other locality data); CMNFI 1979-0789, 1, 164.6 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E); CMNFI 1993-0141, 1, 64.4 mm standard length, Bushehr, Dalaki River (29º28'N, 51º15'E); ID? CMNFI 2007-0061, 2, ? mm standard length, Fars, qanat pool at Ab-e Barik (ca. 27º52'N, ca. 54º09'E); CMNFI 2007-0063, 6, ? mm standard length, Fars, Mand River outside Jahrom (28º36'N, 53º37'E); USNM 205890, 2, 46.0-48.7 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E); ZSM 25705, 1, 107.0 mm standard length, Fars, Lake Parishan (29º45'N, 53º40'E).

Comparative material:- CMNFI 1980-0811, 2, 82.6-112.4 mm, Turkey, Akziyaret Deresi, Tigris River system (no other locality data);

BM(NH) 1931.12.21:1-2, 2, 69.5-78.5 mm standard length, Iraq, Mosul (36º20'N, 43º08'E); BM(NH) 1974.2.22:1184, 1, 130.2 mm standard length, Iraq, Sulaimaniyah (); BM(NH) 1974.2.22:1196, 1, 53.0 mm standard length, Hawiya Canal, Lesser Zab (); BM(NH) 1974.2.22:1214-1255 (in part), Khalis (33º49'N, 44º32'E).

Cyprinion milesi
(Day, 1880)


Head, BMNH 1883.8.2:2-3, Baluchestan, Sib near Dizak

Common names

None.

[sabzug in Pakistan].

Systematics

Barbus milesi was described from "a spring at Tràl", Pakistan.

Berg (1949), Mirza (1969), Mirza et al. (1991) and Howes (1982) recognise this species as valid. If so, synonyms according to Berg (1949), would be Barbus bampurensis Nikol'skii, 1899 described from "Flum. Bampur", Scaphiodon daukesi Zugmayer, 1912 from "Irrigation channels and pools near Panjgur, Baluchistan, Pakistan", and Barbus baschakirdi Holly, 1929 from "Ein Bach bei Guadjik am Wege von Sarzeh in Biabun nach Darpahan in den Bergen von Baschakird, Südostpersien" (= a brook at Guadjik on the way from Sarzeh in Biabun to Darpahan in the Baschakird Mountains, southeast Persia).

Much of my material from southeastern Iran was assigned by me to C. watsoni. Specimens that resemble C. milesi (lacking a shallowly arched or sector mouth with a horny edge but having an oblique u-shaped mouth) are found at the same sample localities as typical C. watsoni. The mouth structure of the putative C. milesi resembles that of juvenile C. watsoni, possibly retained in the adult (paedomorphosis). A Principal Components Analysis does not separate these two forms when the mouth characters are not included in the analysis.

A specimen in the Naturhistorisches Museum Wien under NMW 52736, 34.4 mm standard length, is listed as a syntype under the name Cirrhina milesi but its locality is Gwadur, Hubb River and the type status may be an error.

Five syntypes of Barbus bampurensis, 32.0-64.8 mm standard length, are in the Zoological Institute, St. Petersburg (ZISP 11715) from "Flum. Bampur, 15-23.VII.1898, Zarudnyi". The jar label gives a date of 15-19.VII.1898.

The holotype of Barbus baschakirdi, 52.2 mm standard length, is in the Naturhistorisches Museum Wien under NMW 13798 and a cotype (syntype) of Scaphiodon daukesi, 102.8 mm standard length, is under NMW 19784.

Scaphiodon daukesi types in Munich were destroyed in World War II but one syntype is in the Naturhistorisches Museum Wien under NMW 19784, and two syntypes are in the Zoological Survey of India, Calcutta under ZSI F8028/1 and 8032/1 ((Menon and Yazdani, 1968; Eschmeyer et al., 1996; Neumann, 2006).

Key characters

The mouth is characteristically oblique, longer in lateral view than C. watsoni.

Morphology

The oblique mouth reaches back to the anterior eye margin in small fish and to the rear of the nostril in larger fish. Dorsal fin with 3 unbranched and 10-13 branched rays, anal fin with 2 unbranched and 7 branched rays, pectoral fin with 14-16 branched rays and pelvic fin with 7-8 branched rays. Total gill rakers 11-12. The following description is based mostly on Barbus bampurensis types. Dorsal fin spine strong and serrated, with large teeth in small fish. Lateral line scales 34-39. The scaleless groove before the dorsal fin is weakly expressed. Scales are present on the belly of large fish, almost absent on small fish. Upper flank scales may be regularly or irregularly arranged. Scales have few to no anterior radii, numerous posterior radii, numerous fine circuli, a subcentral anterior focus, and an anterior scale margin indented above and below the mid-line. A pelvic axillary scale is present. The head is more massive in relation to the body than for similar size C. watsoni/kirmanense specimens. The barbel is quite stubby at the base but tapers rapidly to the tip in larger fish. The type series of Barbus bampurensis (= C. milesi) has dorsal fin branched rays 10(4) or 11 (1), anal fin branched rays 7(5), pectoral fin branched rays 14(1) or 15(3) (one unclear), pelvic fin branched rays 7(1) or 8(4), lateral line scales 34(1), 36(1) and 37(3), and total gill rakers 11(3) or 12 (2). Two fish from Sib (see below) had dorsal fin branched rays 9(1) or 10(1), anal fin branched rays 7(2), pectoral fin branched rays 15(2), pelvic fin branched rays 6(1) or 7(1), lateral line scales 35(1) or 37(1), pharyngeal teeth 4,3,2 on the left side, total gill rakers 13(1) or 14 (1), and total vertebrae 38(1) or 39(1). Pharyngeal teeth have a slight hook on the anteriormost tooth with the rest in the main row with scooped-out crowns.

Sexual dimorphism

Tubercles line the anal fin rays and are apparent on the snout in males.

Colour

Copper-brown on the back and upper flank fading to a pinkish belly. Fins are pink and the lateral line has a bright orange streak along it. The preopercle also has orange-golden spots as does the base of the pectoral fins. There is a dark blotch at the base of the caudal fin. The caudal fin base bears a spot in small specimens and there are some much smaller, irregular spots on the caudal peduncle. Peritoneum brown to black.

Size

Attains about 19.0 cm.

Distribution

In Iran, it is recorded from the Sarbaz River of the Makran according to Saadati (1977), the Bampur River of the Hamun-e Jaz Murian basin according to Berg (1949) and the Dozdan River of the Hormuz basin (H. R. Esmaeili). Also in the Mashkid River basin in Pakistan and in rivers draining to the Indian Ocean.

Zoogeography

See under the genus.

Habitat

Unknown.

Age and growth

Unknown.

Food

Unknown.

Reproduction

Unknown.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The distribution, abundance and biology of this species in Iran is poorly known and an assessment for conservation status cannot be given.

Further work

See above.

Sources

Type material: See above, Barbus bampurensis (ZISP 11715),Barbus baschakirdi (NMW 13798) and Scaphiodon daukesi (NMW 19784).

Iranian material: BM(NH)1883.8.2:2-3, 2, 72.2-130.9 mm standard length, Baluchestan, Sib near Dizak (27º15'N, 62º05'E). BWC97-4 no fish on cat sheet?

Comparative material: BM(NH) 1889.2.1:263-264, 2, 89.3-108.7 mm standard length, Afghanistan (no other locality data).

Cyprinion tenuiradius
Heckel, 1847

Common names

botak.

[Araxes kingfish (Fricke et al., 2007)]

Systematics

The type locality is the "Kara-Agatsch als aus dem Araxes" (= Qarah Aqaj River and the Kor River, Fars). Sometimes spelt tenuiradiatus (e.g. in Rainboth (1981) but this is incorrect). Syntypes of Cyprinion tenuiradius are in the Naturhistorisches Museum Wien according to Kähsbauer (1964) under NMW 52808 (1 specimen, 116.7 mm standard length), 52809 (2, 52.3-58.0 mm standard length), 52811 (4, 42.7-47.4 mm standard length), 52815 (1, 77.0 mm standard length) and 52816 (2, 75.5-80.8, although Kähsbauer lists only 1 while Banarescu and Herzig-Straschil (1995) list 2 as also found by me). Other material marked as syntypes from the "Kara-Agatsch. Th. Kotschy" includes NMW 52810 (2 , 103.7-110.0 mm standard length), NMW 52812 (2, 103.5-104.8 mm standard length), NMW 52813 (2, 97.7-103.1 mm standard length), NMW 52814 (1, 114.9 mm standard length), and 52817 (1, not examined). The catalogue in Vienna lists 8 specimens in one column and 26 in the adjacent column. Eschmeyer et al. (1996) add 2 fish from the Araxes River, formerly in NMW, now at the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 2486. The lectotype as selected by F. Krupp in 1984 is NMW 52814 and is published by Banarescu and Herzig-Straschil (1995) with NMW 52808, 52809, 52810, 52811, 52812, 52813, 52815 and 52816 as paralectotypes.

Karaman (1971) assigns this taxon as a subspecies of Cyprinion macrostomum and Bianco and Banarescu (1982) suggest it may be a subspecies in a polytypic species. Berg (1949) records it from the Tigris River where it may be sympatric with C. macrostomum. He considers it to be close to that species, perhaps its southeastern subspecies. Howes (1982) considers tenuiradius to be a variant of C. macrostomum.

Heckel (1847b) distinguishes this species from C. macrostomum by a lower scale count (35-36 as opposed to 42; Berg (1949) gives 35-38 as opposed to 37-43); Krupp (1985c) gives 34-38 compared to 39-43 in macrostomum; Banarescu and Herzig-Straschil (1995) give 36-38, rarely 35 or 39 in C. tenuiradius compared to 41-44, rarely 40 or 45 in C. macrostomum), slenderer body, and a much thinner dorsal spine which is soft in its distal third. The mouth is arched and there is some lower lip development at the mouth corner as in C. kais (see illustrations in Krupp (1985c)). In addition, Berg (1949) gives a branched dorsal fin ray count of 12-13 in C. tenuiradius, 13-15 in C. macrostomum, although Banarescu and Herzig-Straschil (1995) give (12)13-15 for C. tenuiradius from the type locality of Kara-Agasch (sic). Krupp (1985c) states that tenuiradius has a smaller number of scale radii than macrostomum, radii are divergent and the posterior scale margin is curved. However, data for specimens examined by me show overlaps in meristic characters; although means differ, individual fish would be difficult to distinguish on counts alone.

The question then arises as to whether tenuiradius is distinct from macrostomum or merely a variant of a wide-ranging, variable species. The only absolute character is a weaker dorsal fin spine based on examination of type material; other, meristic characters overlap and minor variations in body form are difficult to quantify given a wide range of habitats (lowland rivers and marshes versus highland streams) which may affect shape. The species tenuiradius is retained here as distinct but would benefit from further analyses using new characters, if available, from molecular data.

Key characters

Distribution and a weak spine distinguish this taxon. The dorsal fin spine in macrostomum has teeth extending further along the spine, teeth are more well-developed even near the tip. Spine teeth in tenuiradius are more graded in size as they near the tip and are finer than in macrostomum.

Morphology

Dorsal fin with 4 unbranched and 11-15 branched rays (Berg (1949) has 12-13). The anal fin has 3 unbranched and 6-8 branched rays, usually 7. In 199 Iranian fish, 96.5% have 7 anal fin rays with the rest having 6 rays and 1, presumably anomalous fish, with 9 rays. Pectoral fin branched rays 13-18, pelvic fin branched rays 7-9. Lateral line scales 32-39. Gill rakers 10-21. Scales on the belly may be small and skin covered. There is a naked dorsal keel in front of the dorsal fin, although the area behind the occiput may be scaled and the groove begins nearer the dorsal fin. The mouth is transverse to more or less curved. The dorsal fin spine is weak and serrated only half way or two-thirds of its length. The chromosome number is 2n=50, comprising 13 metacentric, 5 submetacentric and 7 subtelocentric chromosomes pairs. Arm number is NF=86 (Esmaeili and Piravar, 2006).

Meristics for fish from Persian Gulf drainages of Fars, Bushehr and Hormozgan provinces including the Lake Maharlu endorheic basin:- dorsal fin branched rays 11(4), 12(51), 13(175), 14(74) or 15(9) (mean = 13.1, S.D. = 0.746); pectoral fin branched rays 13(3), 14(38), 15(117), 16(41), 17(2) or 18(1)(mean = 15.0, S.D. = 0.733); pelvic fin branched rays 7(23), 8(177) or 9(3)(mean = 7.9, S.D. = 0.345); total gill rakers 10(2), 11(16), 12(27), 13(24), 14(49), 15(35), 16(20), 17(14), 18(8), 19(3) or 21(1)(many counts are based on small specimens and may be low accordingly in comparison with Tigris River basin fishes; mean = 14.2, S.D. = 2.003); and lateral line scales 32(1), 33(15), 34(28), 35(41), 36(47), 37(56), 38(13) or 39(2)(mean = 35.7, S.D. = 1.431).

Sexual dimorphism

Unknown.

Colour

Overall colour is yellowish-white with a light grey back. Scale bases on the flank above the lateral line are brown. The pectoral and pelvic fins have an orange-yellow spot at their base.

Size

Reaches 16.3 cm (Berg, 1949).

Distribution

This species is found in the Gulf and Lake Maharlu basins in Iran (Bianco and Banarescu, 1982; M. Rabbaniha, pers. comm., 1995; Abdoli, 2000; Esmaeili and Gholamifard, 2012).

Heckel's description records this species as from the "Araxes", the modern Kor River in Fars. However, the catalogue sheets in Vienna for the types only list the "Kara Agatsch" (= Mand River) and no subsequent collections have been made of this species in the internal Kor River basin although Abdoli (2000) also maps it from the middle to lower Kor River, possibly based on Heckel's report. Berg (1949) records it from the Tigris River basin, perhaps in error, and Fricke et al. (2007) have it in Turkey from the Aras River system of eastern Turkey (presumably a confusion of the modern Aras or Araxes River with the classical Araxes or Kor River of Fars).

Zoogeography

See under the genus.

Habitat

Unknown in detail but found in springs, streams and rivers of varying descriptions.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 40 fish measuring 5.04-13.49 cm fork length. The a-value was 0.0139 and the b-value 3.063 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Unknown.

Reproduction

Esmaeili and Gholamifard (2012) described the ultrastucture of the chorion and micropyle of the unfertilsied egg.

Parasites and predators

None reported from Iran.

Economic importance

None.

Conservation

The distribution, abundance and biology of this species in Iran is poorly known and an assessment for conservation status cannot be given. Endangered in Turkey (Fricke et al., 2007) but probably does not occur there.

Further work

See above.

Sources

Type material: See above, Cyprinion tenuiradius (NMW 52808, 52809, 52810, 52811, 52812, 52813, 52814, 52815, 52816).

Iranian material: ? see above and ID

Cyprinion watsoni
(Day, 1872)


Ventral head

Common names

None.

[sehrgoar; sabzug = watsoni and microphthalmum - all in Pakistan].

Systematics

Scaphiodon irregularis Day, 1872 described from "rivers in the Sind hills", India, probably Scaphiodon microphthalmus Day, 1880 from "Quetta", Scaphiodon muscatensis Boulenger, 1887 from Muscat, Oman, Cirrhina afghana Günther, 1889 from "Nushki (N. Baluchistan)" and "small river at Kushk (N.W. Afghanistan), Badghis", Cyprinion kirmanense Nikol'skii, 1899 from "Schur-Ab in Kirmano orient.", Cirrhina afghana var. nikolskii Berg, 1905, Scaphiodon macmahoni Regan, 1906, Scaphiodon baluchiorum Jenkins, 1910 (see below for type locality), Scaphiodon watsoni var. belense Zugmayer, 1912 from the "Purali River, near Las Bela" (in Pakistani Baluchistan), Scaphiodon readingi Hora, 1923 from the "Salt Range, Punjab", India, and Cyprinion microphthalmum infraspecies nikolskii Berg, 1949 described originally in part as Cirrhina afghana var. nikolskii Berg, 1905, and Semiplotus dayi Fowler, 1958 are synonyms.

Semiplotus dayi was coined by Fowler to replace Scaphiodon aculeatus, a misidentification by Day (1880) for Chondrostoma aculeatum (= Capoeta aculeata). Fowler thought that Day's fish represented a new species which he named Semiplotus dayi. Howes (1982) considers Semiplotus dayi to be a synonym of Capoeta capoeta (since Karaman (1969a) synonymises Scaphiodon aculeata with C. capoeta. Day's Scaphiodon aculeatus is placed in the synonymy of Cyprinion microphthalmum infraspecies nikolskii by Berg (1949).

Syntypes (or at least specimens examined by Day) of Scaphiodon watsoni described from rivers on the Sind Hills and the Salt Range of the Punjab, India are in the Zoological Survey of India, Calcutta under ZSI 2596 (1), the Natural History Museum, London under BM(NH) 1889.2.1.370-9 (10, but 14 in jar September 2007, 35.6-93.4 mm standard length), the Australian Museum, Sydney under B.7751 (1), the Zoölogisch Museum, Universiteit van Amsterdam under ZMA 115.924 (2) and ZMA 119.255 (1), the Naturhistorisches Museum Wien under NMW 51671 (1), NMW 51672 (1) and NMW 51673 (1), the Museum für Naturkunde, Universität Humboldt, Berlin under ZMB 11042 (1)(132.6 mm standard length), the Rijksmuseum van Natuurlijke Historie, Leiden under RMNH 8704 (1) (or possibly 2552), the Zoological Institute, St. Petersburg under ZISP 8278 (4 but only 2 fish found by me, 63.6-79.6 mm standard length), and the Field Museum of Natural History, Chicago under FMNH 2303 ? 2302(4, 34.0-72.5 mm standard length as examined by me) (Whitehead and Talwar, 1976; Nijssen et al., 1993; Eschmeyer et al., 1996; Ferraris et al., 2000). The 3 fish in the Naturhistorisches Museum Wien measure 86.6, 80.8 and 93.3 mm standard length respectively and are listed there as syntypes.

ZISP 8279 comprising 3 fish, 51.5-52.1 mm standard length, has the same data as ZISP 8278 and may also be types. It is not clear if these are all types, those in ZISP not being marked as types and those in BM(NH) being marked as "possible types"; they may include material simply collected by Francis Day.

A cotype of Scaphiodon watsoni var. belense (NMW 19833) measures 136.9 mm standard length. Eschmeyer et al. (1996) report 2 fish under NMW 19833 although the Vienna card index in 1997 lists only one syntype under this number. In the Zoological Survey of India, Calcutta there are single syntypes under ZSI F827/1 (a misprint for 8027), ZSI F8029/1, ZSI F8030/1 and ZSI F8031/1 (see also Menon and Yazdani (1968)). The remainder of 42 syntypes were in the Munich Museum but were destroyed in World War II (Neumann, 2006).

Types of Scaphiodon microphthalmus are probably lost. The species was described from 2 specimens taken at Quetta in Pakistan. One specimen was sent to the Florence Museum but a recent search failed to locate it and the other specimen has not been located (Whitehead and Talwar, 1976; Banister and Clarke, 1977). A fish measuring 130.1 mm standard length in the Naturhistorisches Museum Wien is listed as a possible syntype (NMW 55897) and in the 1997 card index as "? holotype" (sic).

Note Howes (1982) and Mirza et al. (1991) consider Cyprinion microphthalmum to be a valid species with muscatensis, afghana, afghana var. nikolskii and baluchiorum as synonyms. Howes places macmahoni in watsoni rather than microphthalmum as Berg (1949) and Mirza (1969) do. Howes (1982) also includes irregularis, kirmanense, and readingi in watsoni.

A syntype of Scaphiodon irregularis is in the Australian Museum, Sydney under AMS B.7883 (Ferraris et al., 2000). Syntypes of Scaphiodon muscatensis are in the Natural History Museum, London under BM(NH) 1885.11.7:35-40 (6, 66.4-89.3 mm standard length) and BM(NH) 1887.11.11:289-291 (3, 72.1-79.3 mm standard length) (Eschmeyer et al., 1996; personal observations). Syntypes of Scaphiodon readingi are in the Zoological Survey of India, Calcutta under ZSI F10353/1 and ZSI 10354/1 (27) (sic, although the catalogue numbers seem to indicate only 2 fish) (Menon and Yazdani, 1968) and in the Zoological Museum of Moscow University (ZMMU) (P-1588) (Pavlinov and Borissenko, 2001).

Three syntypes of Scaphiodon baluchiorum (ZSI F9398 to F9400) and one syntype of Scaphiodon macmahoni (ZSI F1239/1) are in the Zoological Survey of India, Calcutta (Menon and Yazdani, 1968). A syntype of Scaphiodon macmahoni measuring 58.6 mm standard length from "Seistan" is in the Natural History Museum, London and was labelled as Cyprinion watsoni (BM(NH) 1905.11.29:27, 58.6 mm standard length). The type locality of Scaphiodon baluchiorum is "Gishtigan (Bampusht); Kalagan, 3,500 feet; Baluchistan". These localities are in Pakistani Baluchistan; Gishtigan being on the Kulushta River which drains into the Nihing River and then the Dashti River (Jenkins, 1910) (these are near the border of Iranian Baluchestan with the upper reaches of the Nihing being in Iran) and Kalagan possibly being the Kalugar River with headwaters in Iran and draining to the Hamun-i Mashkel in Pakistan. The type locality of Scaphiodon macmahoni is "affluents of the Helmand" (Regan, 1906), presumably an error for "effluents" or the delta of the Helmand.

The holotype of Cyprinion kirmanense, 61.6 mm standard length, is in the Zoological Institute, St. Petersburg under ZISP 11712 from "Schur-Ab in Kirmano orient. 27.VI." The 5 syntypes of Cirrhina afghana var. nikolskii are in the Zoological Institute, St. Petersburg (ZISP 11709) and are from the "Bampur River, 27 VII 1898, N. Zarudnyi" according to Berg (1949) but he mentions 2 additional fish with a somewhat deeper body, presumably also part of the type series. ZISP 11709 does have 7 specimens, 43.0-79.1 mm standard length, with a date 15-27.VII.1898. Four syntypes of Cirrhina afghana measuring 74.6-83.0 mm standard length from "Kushk" annotated Afghan. Boundary Comm. are in the Natural History Museum, London (BM(NH) 1886.9.21:150-154 - note that 150-154 indicates there should be 5 fish) with a further 6 syntypes measuring 44.9-99.5 mm standard length labelled "Nushki" and also annotated Afghan. Boundary Comm. (BM(NH) 1886.9.21:155-159 - note this indicates there should be 5 fish in this jar and probably one fish has been mixed up). Additional syntypes are in the Zoological Survey of India, Calcutta under ZSI 11474-11476 (3) and ZSI 11479-11485 (7) (Eschmeyer et al., 1996).

Berg (1949) places Cirrhina afghana var. nikolskii in his Cyprinion microphthalmum infraspecies nikolskii (see also Berg (1933a)). This infraspecies occurs together with Cyprinion microphthalmum but differs by a stronger osseous ray in the dorsal fin which is serrated almost to the summit (Berg (1949) states that transitions exist). The anterior belly region is scaleless also. ZISP 11709 fish mostly have their dorsal spines snapped off but one fish has osseous ray teeth between three-quarters and four-fifths along the spine and a second about three-quarters. ZISP 25406 from a qanat between Kerman and Bandar-e `Abbas comprises 12 fish, 31.0-53.6 mm standard length, belonging to infraspecies nikolskii according to Berg (1949). These fish, of all sizes, have the last quarter to a third of the osseous spine in the dorsal fin unserrated. The mouth form varies. One large fish has a terminal mouth, moderately oblique in lateral view, and no strong horny layer on the lower jaw. Others have a u-shaped or horny jaw positioned on the lower head surface so there is no real gape in lateral view. Some small fish are transitional between the two types. Fin serration, mouth form and development of scales on the anterior belly seem to be widely variable within samples of Cyprinion from a single locality and presumably a single species.

Berg (1949) recognises Cyprinion watsoni belense as a subspecies, rather than a variety as originally described, from Indian Ocean drainages of southeastern Iran and southwestern Pakistan (Baluchistan). It is distinguished by smaller scales (33-36) from the type form (31-34), hardly a sufficient criterion given the wide distribution range and individual variation shown by these fishes.

This species has not been adequately examined in southeastern Iran and most nominal species are referred to Cyprinion watsoni, the earliest available name for the taxon. C. watsoni is distinguished from other Iranian Cyprinion by having 9-11 dorsal fin branched rays (macrostomum and tenuiradius have 12-15; C. milesi also has a low dorsal ray count but has an oblique mouth, not transverse or arched (Berg, 1949)). Bianco and Banarescu (1982) consider that several subspecies may eventually be defined and that some of the names in synonymy here would then be used.

Berg (1949) also recognises C. irregulare as a distinct species with a low dorsal fin branched ray count as in C. watsoni but usually 37 or more scales in the lateral line, a scaleless groove on the back before the dorsal fin, and upper scale rows anteriorly arranged irregularly and not imbricate and C. microphthalmum with a low dorsal fin branched ray count as in C. watsoni but usually 37 or more scales in the lateral line, a scaleless groove on the back before the dorsal fin barely outlined, and upper scale rows anteriorly arranged regularly and imbricate. C. microphthalmum infraspecies nikolskii is described as having a strong dorsal fin spine with obvious teeth extending to the tip while typical C. microphthalmum has a weak ray with weak teeth only visible when the skin covering the fin is peeled away.

Berg (1949) later states that no great importance should be attached to the upper row scale arrangement and the groove development - if the groove is well-developed then the upper row scales are irregular and this phenomenon can be seen in some C. watsoni and C. microphthalmum specimens. Berg then suggests that irregulare could be regarded as an infraspecies of C. microphthalmum as this type of condition occurs in Capoeta fusca and in Garra rossica. Under the heading C. watsoni Berg also gives mouth shape, scale arrangement, dorsal fin spine serrations, and body form as characters which can vary greatly. These observations serve to confirm the great variability in characters for these fishes. Large series of adults and young would be needed to adequately define species and subspecies.

Mirza (1969) reports C. watsoni, C. microphthalmum and C. milesi from western Pakistan and Iran, the former in Makran drainages and the latter two in the Mashkel (= Mashkid) River basin. The characters used to separate these taxa are an oblique mouth and head length contained less than 4.5 times in total length (= C. milesi), an arched mouth, head length more than 4.5 times in total length, scaleless strip on back conspicuous, and 33-36 lateral line scales (= C. watsoni), and a transverse mouth, head length more than 4.5 times in total length, scaleless strip on back hardly visible, and 37-40 lateral line scales (= C. microphthalmum). Sample sizes in this study were small (22 fish) and these characters show considerable variation in larger samples and between fish of different sizes.

Key characters

The arched mouth and and 9-12, usually 10-11, branched dorsal fin rays serve to identify this species.

Morphology

Young fish have a more horseshoe-shaped mouth than larger and older fish where the mouth is a shallow arch, almost straight. The dorsal fin has 3-4 unbranched and 9-12 branched rays, the last unbranched ray of the dorsal fin being variably serrated and thickened. The extent of serrations appears to vary independently of size, from half to three-quarters or more of the spine length. The anal fin has 1-3, usually 3, unbranched and 6-8, usually 7 branched rays. In Iranian specimens, 89.7% of 419 fish had 7 branched anal fin rays, the remainder having 6 branched rays. Pectoral fin branched rays 11-18, usually 15-16, and pelvic fin branched rays 6-9, usually 8. Lateral line scales 31-43. Scales have well-developed anterior radii as well as posterior and some lateral radii. The scale focus is almost central on mid-flank scales. There is a naked median strip on the back in front of the dorsal fin, about one scale wide, in some fish. Some fish may show poor imbrication of scales on the belly and upper anterior flank. Gill rakers 8-18, reaching to or past the adjacent raker when appressed. Total vertebrae 38. Rarely with a tripartite swimbladder, usually bipartite (Mirza, 1971 - for his C. microphthalmum). Pharyngeal teeth 2,3,4-4,3,2 or 2,3,5-5,3,2, with spoon-shaped crowns. The back in front of the dorsal fin is naked in the mid-line. Mouth shape variable, from and arch to a transverse cutting edge. Scales on the belly can be embedded in the skin, obvious or even absent.

Meristics for Iranian specimens:-
 

Locality/Dorsal Fin Rays

9

10

11

12

x

S.D.

Hamun-e Mashkid

16

3

10.2

0.375

Hamun-e Jaz Murian

3

15

5

10.1

0.596

Dasht-e Lut

2

50

4

10.0

0.328

Makran

1

29

3

10.1

0.348

Hormuz

2

144

124

7

10.5

0.562

Sirjan

1

3

7

10.5

0.688

 

Locality/Pelvic Fin Rays

6

7

8

9

x

S.D.

Hamun-e Mashkid

6

13

7.7

0.478

Hamun-e Jaz Murian

1

21

1

8.0

0.302

Dasht-e Lut

1

54

8.0

0.135

Makran

33

8.0

0.000

Hormuz

2

31

237

8

7.9

0.400

Sirjan

2

9

7.8

0.405

 

Locality/Pectoral Fin Rays

11

13

14

15

16

17

18

x

S.D.

Hamun-e Mashkid

9

10

15.5

0.513

Hamun-e Jaz Murian

1

8

11

2

1

15.7

0.864

Dasht-e Lut

1

2

14

31

8

14.8

0.879

Makran

1

14

15

3

15.6

0.704

Hormuz

1

46

111

99

20

1

15.3

0.863

Sirjan

4

4

3

14.9

0.831

 

Locality/Total Gill Rakers

8

9

10

11

12

13

14

15

16

17

18

x

S.D.

Hamun-e Mashkid

1

4

3

4

3

4

13.8

1.718

Hamun-e Jaz Murian

3

4

10

4

1

1

12.0

1.492

Dasht-e Lut

1

7

8

16

12

10

2

12.2

1.427

Makran

1

6

9

4

8

3

1

1

13.0

1.794

Hormuz

3

7

33

45

64

60

41

10

2

1

12.2

1.569

Sirjan

2

2

1

5

1

12.1

1.375

 

 
Locality/Lateral Line Scales 33 34 35 36 37 38 39 40 41 42 43 x S.D.
Hamun-e Mashkid 1 3 9 1 1 1 2 1
Hamun-e Jaz Murian 1 9 11 1 1
Dasht-e Lut 5 12 26 10 2 1
Makran 2 1 17 8 4 1
Hormuz 1 2 17 57 99 72 20 7 3
Sirjan 3 4 3 1

? vertebrae

The holotype of Cyprinion kirmanense has 10 dorsal fin branched rays, 7 anal fin branched rays, 15 pectoral fin branched rays, 8 pelvic fin branched rays, 37 lateral line scales and 13 total gill rakers (not in above tables).

Sexual dimorphism

Males have snout tubercles and tubercles on the anal fin rays (Regan, 1906; Jenkins, 1910; Berg, 1949). Large tubercles are found on the snout in front of the nostrils, the top of the head, and in rows on the rays of the caudal and anal fin, following the ray branching, in a fish not yet fully mature (40.1 mm standard length). ZMB 11042 (132.6 mm SL) has tubercles thickly present on the snout extending back to the nostrils and then to the eyes, scattered all over the sides of the head, absent on top of the head (may be lost in this old specimen) and large tubercles on anal fin rays near the tip. There is a depression in front of the nostrils in adult males.

Colour

The back and upper flank are dark or copper brown, golden or dark olive, light green-brown or brown-grey, sometimes with bluish or orange tinges, fading to a light or yellowish pink on the lower flanks and belly. The flank scales are silvery and may be outlined in black. Black spots may be present along the flank as may be an orange stripe or a series of 7-9 orange spots above the lateral line anteriorly. The orange colour may be deep, almost red. Occasionally, there may be an orange spot below the lateral line. There may be a vertical orange line over the cleithrum or a spot at its postero-ventral corner. In some fish the whole cleithrum area is red-orange. The operculum, preoperculum and cheek can be iridescent blue. The bases of the pectoral and pelvic fins and part of the operculum may also be pink or orange-coloured. Young have a fine black streak above the lateral line. The dorsal and caudal fins are lead-coloured to black and other fins are pink to yellowish. All the fins except the paired fins may be hyaline. There can be a black caudal base spot, quite marked in some fish, particularly small ones. The peritoneum is black or dark brown.

Size

Attains 23 cm (Zugmayer, 1912).

Distribution

From southeastern Iran east to India. In Iran, it is recorded from the Dasht-e Lut, Hormuz, Hamun-e Jaz Murian, Sistan, Hamun-e Mashkid basins and in the Makran from the Jagin to the Bahu Kalat rivers (Berg, 1949; Spillman, 1972; Bianco and Banarescu, 1982; Abdoli, 2000; Esmaeili et al., 2011?).

Zoogeography

The occurrence of this species (as C. microphthalmum in Banister and Clarke (1977)) in Oman across the Straits of Hormuz is a result of the 120 m lowered sea level from 100,000 B.P. to 10,000 B.P. The Tigris-Euphrates then ran down the Gulf and presumably provided ready access across it. Banister and Clarke (1977) comment that it is surprising that only one species made the crossing but nothing is known of the climate during this 90,000 year period nor of the composition of the Iranian fish fauna. It may well have been quite impoverished. Hora (1956) describes fish paintings on pots from Nal in Pakistani Baluchistan dating from the third millenium B.C. One of the species not clearly represented is a Cyprinion, now found in that area (not extinct as Banister (1980) would have it). This potentially shows how the ichthyofauna in Southwest Asia can change over relatively short periods of time; the changes over 100,000 years must have been considerable and not readily traceable.

Dorsal fin branched rays have moderately strong modes of 10 for the Hamun-e Mashkid, Hamun-e Jaz Murian, Dasht-e Lut and Makran basins but modes of 10 and/or 11 for Hormuz and Sirjan, the westernmost basins. C. watsoni may be showing some introgression with Cyprinion species to the west which have higher counts. However, subsamples within the Hormuz basin do not show a clear pattern of higher modes or means towards the west.

Habitat

Kiabi and Abdoli (2000) found this species to be the commonest and to have the widest range in Hormozgan Province.

Age and growth

Unknown. Esmaeili and Ebrahimi (2006) give a significant length-weight relationship based on 23 Iranian fish measuring 8.34-13.38 cm total length. The a-value was 0.0101 and the b-value 2.952 (a b-value < 3 indicating a fish that becomes less rotund as length increases and a b-value >3 indicating a fish that becomes more rotund as length increases).

Food

Gut contents are primarily herbivorous items including filamentous algae such as Cladophora and Spirogyra, and a wide range of diatoms but some insect material is also found (Mirza, 1969; Farooq et al., 1996).

Reproduction

Spawning takes place in Pakistan at Islamabad from mid- to late March to mid-April (Shaikh and Jalali, 1989, 1991) and near Islamabad (33.3°N, 73.0°E) in April and May (Shaikh and Hafeez, 1993). Gonads begin to develop in December as photoperiod and temperature rise but a continuing warm temperature is the predominant factor for spawning to occur; a fall in temperature halts spawning. Eggs are dark yellow when mature, testes creamy when ripe. Spawning occurs once a year. Up to 150 eggs are recorded in fish from the Ab Garm-e Ganow with a diameter of 1.2 mm.

Parasites and predators

Males were reported as having snout tubercles and tubercles on the anal fin rays (Regan, 1906) but these were the encysted glochidia of a unionid mollusc (B. Prashad in Annandale and Hora (1920). Jalali et al. (1995) describe a new species of monogenean, Dactylogyrus pallicirrus, from fish taken in the Shur River, a Halil River tributary in the Hamun-e Jaz Murian basin. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record Gyrodactylus sp. for fish from the Minab and Halil rivers.

Economic importance

This species is of no economic importance although Butt (1995) suggests that it could be a food source in Pakistan, occurring in shoals of considerable size in rivers that otherwise support little in the form of aquatic protein. It could be cultured as food and as a forage fish. This species has been used to study the effects of heavy metals in Pakistan (Shah, 2002). Higher concentrations of copper and zinc caused lethargy and loss of equilibrium.

Conservation

This species is widely distributed in various basins in southeastern Iran and neighbouring areas and does not appear to under any threat.

Further work

The biology of this species requires study and a thorough review of morphology in relation to named taxa should be carried out.

Sources

Type material: See above and note reservations on type status of some, Cirrhina afghana (BM(NH) 1886.9.21:150-154, BM(NH) 1886.9.21:155-159); Cirrhina afghana var. nikolskii (ZISP 11709); Cyprinion kirmanense (ZISP 11712); Scaphiodon macmahoni (BM(NH) 1905.11.29:27); Scaphiodon microphthalmus (NMW 55897); Scaphiodon muscatensis (BM(NH) 1885.11.7:35-40, BM(NH) 1887.11.11:289-291); Scaphiodon watsoni (FMNH 2303, NMW 51671, NMW 51672 and NMW 51673, ZMB 11042, ZISP 8278, ZISP 8279); Scaphiodon watsoni var. belense (NMW 19833).

Iranian material: CMNFI 1979-0138, 49, ?-66.8 mm standard length, Fars-Hormozgan border, stream in Rasul River drainage (ca. 27º32'N, ca. 54º58'30"E); CMNFI 1979-0143, 22, ? mm standard length, Hormozgan, marsh in Hasan Langi River drainage (27º21'N, 56º50'30"E); CMNFI 1979-0144, 76, ? mm standard length, Hormozgan, Minab River at Minab (27º09'30"N, 57º04'E); CMNFI 1979-0145, 139, ? mm standard length, Hormozgan, Geru River south of Minab (26º55'N, 57º01'30"E); CMNFI 1979-0149, 54, ?-93.8 mm standard length, Hormozgan, stream north of Bandar Abbas (27º36'N, 56º14'E); CMNFI 1979-0150, 34, 43.8-101.6 mm standard length, Hormozgan, stream at Gohreh (27º45'N, 56º05'E); CMNFI 1979-0152, 10, ? mm standard length, Homozgan, Shur River drainage (28º09'N, 55º43'E); CMNFI 1979-0153, 31 24.5-84.7 mm standard length, Fars, qanat stream and pool at Qaleh-ye Biabani (28º31'N, 54º53'E); CMNFI 1979-0154B, 49, ? mm standard length, Fars, stream channels at Koorsiah (28º45'30"N, 54º24'E); CMNFI 1979-0155, 26, ? mm standard length, Fars, springa t gavanoo (28º47'N, 54º22'E); CMNFI 1979-0156, 12, ? mm standard length, Fars, qanat at Rashidabad (28º47'N, 54º18'E); CMNFI 1979-0167, 25, ? mm standard length, Kerman, qanat at Bam (29º06'N, 58º20'E); CMNFI 1979-0168, 50, ?-93.8 mm standard length, Kerman, qanat at Shahabad (29º07'N, 58º16'E); CMNFI 1979-0173, 15, 26.5-84.0 mm standard length, Hormozgan, qanat at Hajjiabad (28º19'N, 55º54'E); CMNFI 1979-0176, 1, ? mm standard length, Hormozgan, Sarzeh River (27º30'30"N, 56º15'30"E); CMNFI 1979-0180, 5, ? mm standard length, Hormozgan, stream 3 km east of Essin (27º19'N, 56º17'30"E); CMNFI 1979-0181, 19, ? mm standard length, Hormozgan, Kul River (27º17'30"N, 56º03'30"E); CMNFI 1979-0183, 14, ? mm standard length, Hormozgan, stream in Rasul River drainage (27º11'30"N, 55º42'E); CMNFI 1979-0185, 4, ? mm standard length, Hormozgan, stream in Rasul River drainage (27º06'N, 55º45'E); CMNFI 1979-0186, 11, ? mm standard length, Hormozgan, strean and pools at Sar Khun oasis (ca. 27º24'30"N, ca. 56º25'E); CMNFI 1979-0187, 54, 18.9-73.6 mm standard length, Hormozgan, stream and pools at Sar Khun oasis (27º23'30"N, 56º26'E); CMNFI 1979-0188, 18, ? mm standard length, Hormozgan, jube at Gohreh (27º45'N, 56º05'E); CMNFI 1979-0189, 25, ? mm standard length, Hormozgan, jube and pool on road to Darab (27º08'30"N, 55º42'E); CMNFI 1979-0190, 44, ?-83.3 mm standard length, Fars-Hormozgan border, spring and pool at Galah Tuyeh (ca. 28º32'N, ca. 55º14'E); CMNFI 1979-0191, 35, 36.6-86.5 mm standard length, Fars, stream 10 km east of Furg (ca. 28º16'N, ca. 55º18'E); CMNFI 1979-0219, 19, 19.1-33.0 mm standard length, Kerman, jube 14 km west of Jiroft (28º37'N, 57º41'E); CMNFI 1979-0220, 4, 28.0-65.5 mm standard length, Kerman, jube 2 km south of Jiroft (28º39'N, 57º43'E); CMNFI 1979-0309, 2, 101.2-108.5 mm standard length, Kerman, Fahraj River at Azizabad (28º57'N, 58º42'E); CMNFI 1979-0310, 1, ? mm standard length, Baluchestan, qanat at Bazman (27º49'N, 60º12'E); CMNFI 1979-0311, 10, ? mm standard length, Baluchestan, Bampur River at Malakabad (27º11'N, 60º27'E); CMNFI 1979-0312, 39, ? mm standard length, Baluchestan, dam on Bampur River (27º11'N, 60º36'E); CMNFI 1979-0313, 68, ? mm standard length, Baluchestan, Bampur River at Bangharabad (27º20'N, 60º46'E); CMNFI 1979-0314, 10, 25.5-118.4 mm standard length, Baluchestan, qanat at Karavandar (27º50'N, 60º46'E); CMNFI 1979-0315, 71, ? mm standard length, Baluchestan, Bampur River 2 km north of Karavandar (27º51'N, 60º46'E); CMNFI 1979-0316, 22, 14.5-69.8 mm standard length, Baluchestan, stream 68 km south of Iranshahr (26º48'N, 61º02'E); CMNFI 1979-0317, 11, 16.5-118.6 mm standard length, Baluchestan, Sarbaz River at Bondan (26º35'N, 61º13'E); CMNFI 1979-0318, 11, ? mm standard length, Baluchestan, Sarbaz River at Huvar (26º09'N, 61º27'E); CMNFI 1979-0323, 6, ? mm standard length, Baluchestan, Sarbaz River (ca. 26º26'N, ca. 61º16'E); CMNFI 1984-0324, 4, ?39, ?51.9-117.8 mm standard length, Baluchestan, Bampur River at Sa'idabad (27º11'N, 60º22'E); CMNFI 1979-0325, 7, ? mm standard length, Baluchestan, qanat at Espakeh (26º51'N, 60º14'E); CMNFI 1979-0326, 10, ? mm standard length, Baluchestan, stream south of Pip (ca. 26º35'N, ca. 60º02'E); CMNFI 1979-0327, 10, 24.0-62.4 mm standard length, Baluchestan, stream in Geh River drainage (26º32'N, 59º57'E); CMNFI 1979-0329, 82, ? mm standard length, Baluchestan, stream at Zaminbandan (27º02'N, 61º20'E); CMNFI 1979-0331, 25, 13.1-50.3 mm standard length, Baluchestan, qanat in Saravan (27º22'N, 62º20'E); CMNFI 1979-0332, 9, 20.8-33.3 mm standard length, Baluchestan, qanat at Kalapurkan (27º14'N, 62º33'E); CMNFI 1979-0334, 4, 26.3-38.4 mm standard length, Baluchestan, Mashkid River (27º04'N, 62º54'E); CMNFI 1979-0335, 2, 66.8-72.2 mm standard length, Baluchestan, qanat at Esfandak (27º07'N, 62º50'E); CMNFI 1979-0338, 17, ? mm standard length, Baluchestan, Tahlab River drainage 8 km from Mirjaveh (28º58'N, 61º24'E); CMNFI 1979-0339, 24, 24.5-76.9 mm standard length, Baluchestan, Tahlab River drainage 16 km from Mirjaveh (28º56'30"N, 61º21'E); CMNFI 1979-0411, 8, ? mm standard length, Hormozgan, Minab River (27º24'N, 57º12'E); CMNFI 1979-0412, 22, 22.0-122.2 mm standard length, Hormozzgan, spring at Saras (27º30'N, 57º34'E); CMNFI 1979-0415, 5, ? mm standard length, Hormozgan, stream south of Ab Garm-e Ganow (27º17'30"N, 56º20'E); CMNFI 1979-0416, 2, 40.1-55.9 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E); CMNFI 1979-0418, 5, 58.2-111.2 mm standard length, Hormozgan, river near Kahkom (28º09'N, 55º43'E); CMNFI 1991-0141, 3, ?, mm standard length, ; see original sheet? CMNFI 2007-0031, 12, ? mm standard length, Baluchestan, headwater of Bampur River (27º51'N, 60º46'E); CMNFI 2007-0033, 15, ? mm standard length, Baluchestan, Rusgay qanat in Iranshahr (27º13'N, 60º41'E); CMNFI 2007-0034, 3, ? mm standard length, Baluchestan, headwater stream on road to Zaboli (ca. 26º58'N, ca. 61º27'E); CMNFI 2007-0036, 8, ? mm standard length, Baluchestan, qanat at Bazman (27º49'N, 60º12'E); CMNFI 2007-0037, 7, 62.4-166.3 mm standard length, Kerman, Hosseinabad and Gamatabad qanats at Bam (29º06'N, 58º21'E); CMNFI 2007-0038, 9, 62.8-101.2 mm standard length, Kerman, Mehtiabad qanat at Bam (29º06'N, 58º21'E); CMNFI ?, 11, 46.5-69.1 mm standard length, ; 44 ? check, Sirjan on data sheet but no fish at this number in catalogue, check shelf CMNFI 2007-0049, 11, ? mm standard length, Hormozgan, ditches in upper Kol River basin at Hajjiabad (ca, 28º19'N, ca. 55º55'E); 45 CMNFI 2007-0050, 4, 61.2-92.4 mm standard length, Hormozgan, ditches in upper Kol River basin at Hajjiabad (ca. 28º19'N, ca. 55º55'E); 46 CMNFI 2007-0051, 7, 54.6-84.3 mm standard length, Hormozgan, upper kol River basin at Hajjiabad (28º19'N, 55º55'E); 47 CMNFI 2007-0052, 2, 70.7-92.3 mm standard length, Hormozgan, ditch at Qotbabad (27º46'N, 56º06'E); 48 CMNFI 2007-0055, 15, 24.5-75.3 mm standard length, Hormozgan, headwtaer stream in Minab River basin (ca. 27º47'N, ca. 57º12'E); 51 CMNFI 2007-0056, 14, 30.2-70.4 mm standard length, Kerman, qanat at Kahnuj (27º58'N, 57º45'E); 52 CMNFI 2007-0059, 9, ? mm standard length, Fars, Chashmeh Barashk (ca. 27º24'N, ca. 54º06'E); CMNFI 2007-0060, 3, 56.2-93.7 mm standard length, Fars, Chashmeh Ab-e Shirin near Lar (ca. 27º41'N, ca. 54º17'E); 57ID? BM(NH) 1883.8.2:4-9, 5, 42.4-114.9 mm standard length, Baluchestan, Jalq (27º36'N, 62º42'E); BM(NH) 1883.8.2:20-25, 6, 23.1-85.2 mm standard length, Baluchestan, Sib near Dizak (27º15'N, 62º05'E); NMW uncatalogued, 19, 18.1-52.9 mm standard length, Hormozgan, Ab Garm-e Ganow (ca. 27º26'N, ca. 56º20'E). BWC97-4, ?, ? mm standard length, Kerman, Halil River (no other locality data); no fish on cat sheet? BWC97-5, 1, ? mm standard length, Hormozgan, Hasan Langi River (no other locality data);

Comparative material: CAS 28722, 1, 117.4 mm standard length, India, Punjab, Salt Range, Katas Nallah (no other locality data);

Genus Cyprinus
Linnaeus, 1758

The carp genus is found in Europe and Asia and comprises several species of which one has been widely introduced as a food fish.

This genus is characterised by a compressed but heavy body, large size, rounded snout, 2 pairs of barbels, large molar pharyngeal teeth in 3 rows, a very long dorsal fin with the last unbranched ray spine-like and serrated, the anal fin short but with the last unbranched ray spine-like and serrated, the gut is moderately long, and the dorsal and lateral skull bones are sculptured.

Cyprinus carpio
Linnaeus, 1758

 


Caught on sweet corn bait fished on bottom in Anzali Mordab near Abkenar Village,
15 June 2012, 37 cm, 1.1 kg, courtesy of S.Nouripanah

Duane Raver, U.S. Fish and Wildlife Service
Duane Raver, U.S. Fish and Wildlife Service

Mirror carp, Duane Raver, U.S. Fish and Wildlife Service
Mirror carp, Duane Raver, U.S. Fish and Wildlife Service

Common names

كپور (= kopur, kapur or kapoor (in Gilaki)), kapur-e ainehi (= mirror carp), كپور معمولي (kapur-e ma'mouli or mar'mulleh or ma'muli or maamoli (= common carp)), rashti or كپور رشتي (kapur-e Rashti) in Khuzestan (because their origin was Rasht), mahi-ye gul (= flower fish, meaning in this sense a good fish).

[car or carp shaeeh in Arabic; caki in Azerbaijan; geitan-tsatsan or dliter, both in Armenia; sazan for wild and karp for cultured carp in Russian; common carp, European carp, German carp, wild carp; mirror carp, leather carp, line carp, naked carp (last four referring to scalation), koi (aquarium variety)].

Systematics

Cyprinus Carpio was originally described from Europe. Cyprinus carpio var. caspicus Walbaum, 1792 is described from the mouth of the Volga and Don rivers but is infrasubspecific and the name has not been used in Iran nor has Cyprinus carpio fluviatilis Pravdin, 1945 described from floodplain lakes of the Volga River near Saratov. Mousavi Gel Sefid et al. (2007) give some morphometric and meristic characters of fish from the Anzali Lagoon. Ghelichipour et al. (2011) examined fish from the Qarasu and Anzali regions of the Iranian Caspian shore using microsatellite markers and found two different populations.

Key characters

This species is easily identified by the long dorsal fin, the spine in both the dorsal and anal fins, and the two pairs of barbels.

Morphology

Dorsal fin with 2-5 unbranched rays followed by 14-23, usually 18-20, branched rays, anal fin with 2-4, usually 3, unbranched rays followed by 3-7, usually 5, branched rays, pectoral fin branched rays 13-19, and pelvic fin branched rays 5-9, usually 8. The dorsal fin has the last unbranched ray developed as a toothed spine and the anal fin has a similar spine. Lateral line scales 26-41, mostly 36-39. Scales may be absent (leather carp), restricted to a few, enlarged scales (mirror carp), or only a mid-lateral row of scales (line carp), in cultivated varieties. Wild carp are fully scaled. Individual scales have a central focus, wavy anterior margin, few radii on the anterior and posterior fields in young fish, and medium numbers of radii on fish 12-14 cm standard length. There are numerous fine circuli and the posterior scale field breaks up into bubble-like structures. Gill rakers 17-29 (some literature counts may be lower arm of arch only and there may be size-related variation too) and vertebrae 32-39 (lower counts may not include Weberian vertebrae). Rakers touch the second raker below when appressed and have a row of knobs on their medial surface. Pharyngeal teeth 1,1,3-3,1,1 with variants 1,2,3-3,2,1, 1,2,3-3,1,1, 1,1,1,3-3,1,1, 1,1,3-2,1,1 and 1,3-3,1,1. Posterior major row teeth are large with flattened crowns bearing wavy ridges while more anterior teeth are a rounded knob, or even concave on top of the knob. The gut is elongate with several coils. This species is a tetraploid (2n=98-104)(Al-Sabti, 1986; Klinkhardt et al., 1995).

Meristic values for Iranian specimens are:- dorsal fin branched rays 16(1), 17(5), 18(18), 19(16), 20(10); anal fin branched rays 4(1) or 5(49); pectoral fin branched rays 14(1), 15(7), 16(25), 17(16) or 18(1); pelvic fin branched rays 6(1), 7(6) or 8(43); lateral line scales 34(2), 35(6), 36(10), 37(26) or 38(6); total gill rakers 18(1), 20(2), 21(11), 22(11), 23(15), 24(4), 25(4) or 26(2); total vertebrae 36(1), 37(19), 38(26) and 39(6); and pharyngeal teeth ?. Note that these samples may include individuals which are not native but introduced for fish farming from stocks outside Iran.

There are morphometric and meristic differences between carp from the southwestern and southeastern Caspian Sea but these are ecological not taxonomic. Also, carp from the Anzali Mordab differ from those in the sea by having a longer head, snout, eye and postorbital region (although of course some of these are redundant), greater interorbital width, shorter predorsal and preanal distances, shorter dorsal, anal, pectoral and caudal fins, and a lower anal fin (A. M. Shukolyukov in Berg (1948-1949)). Yousefian (2004) found carp from the Caspian Sea in Iran had a dominant genotype different from those in a fish farm. Meristics (scales and fin rays) and morphometrics (head length and body width) also differed.

Sexual dimorphism

Females are deeper bodied than males because of their eggs and the distance between the pectoral and pelvic fins and the pelvic and anal fins is more. Dorsal and anal fins in males are higher, the anal fin is longer at the base, the pectoral fin is longer and the lobes of the caudal fin are longer. This is accounted for by the greater swimming activity of males during spawning (Kuliyev and Agayarova, 1984). Breeding males have fine tubercles on the head, particularly on the anterior operculum and preoperculum and under the eye, above the lateral line and more frequently below it, and on the fin rays.

Colour

In semi-diadromous carp from the Kura region of the southwest Caspian Sea, the overall body colour is dark yellow, the flanks being golden-yellow with dark shading, and the back is black. The belly and fins are light yellow and the caudal fin is reddish. Lake populations are darker. Young fish from Iran are silvery on the flanks (but not as bright as Carassius auratus), greyish on the back, silver-pearl on the belly, the iris is silvery with grey above and below, the dorsal fin and upper caudal lobe are pale grey, the lower caudal lobe and anal fin are orange, the pelvic fin is pale orange, and the pectoral fin has only traces of orange. The caudal fin may be yellow-orange with lobe margins red. The freshwater resident form in the Anzali Mordab is yellowish, the semi-diadromous form dark. The peritoneum is grey to silvery and may be speckled.

Size

Carp resident in fresh waters are smaller than semi-diadromous carp. In the 1950s in Iran, carp catches were 20-41 cm long (Farid-Pak, no date). Fish up to 1.0 m long are caught in the Caspian basin (A. Abdoli, pers. comm., 1995). Maximum size reaches 1.28 m and 46.1 kg, possibly to 1.5 m and 69.6 kg (Machacek (1983-2012), downloaded 27 July 2012).

Distribution

Widely introduced in the Middle East for aquaculture. Found naturally in Iran in the whole Caspian Sea drainage, it is also widely stocked in the provinces of Gilan and Mazandaran (Petr, 1987; Abbasi et al., 1999). It is reported from the Atrak, Gorgan, Gharasu, Tajan, Babol, Aras, and Safid rivers, the Anzali Mordab and the Siah-Keshim Protected Region, Boojagh Wetland, Gorgan Bay, the southeast Caspian Sea, southwest Caspian Sea and south-central Caspian Sea as an introduced species, and in all these plus the Haraz and Pol-e Rud rivers as a native species (Riazi, 1996; Kiabi et al., 1999; Khara et al., 2011).

It is probably native to the Tedzhen River of Turkmenistan (the Hari River or Harirud in Iran) (Aliev et al., 1988). This species is also recorded from the Karakum Canal and Kopetdag Reservoir in Turkmenistan (Shakirova and Sukhanova, 1994; Sal'nikov, 1995) and may eventually reach Iranian waters in the Tedzhen (= Hari) River basin. These latter stocks may be introduced.

It has been introduced to the Sistan basin to a canal flowing into the Chahnimeh, the Sistan Dam, Hamun Lake and the Hamun Kushk (Ahmadi and Wossughi, 1988; J. Holčík, in litt., 1996; Shamsi et al., 2009)), to the Karaj River in the Namak Lake basin, to the Mahabad Dam (Abdi, 1999; www.mondialvet99.com, downloaded 31 May 2000), to the Haft Barm lakes near Shiraz in 1984 (Petr, 1987), to the Manjil Reservoir on the Safid Rud (Nümann, 1966), to Mehr Dasht Lake, Markazi, to the Zayandeh River Dam (Y. Keivany, in litt., 1992; Ghorbani Chafi, 2000), to Lake Zaribar, Kordestan (Abzeeyan, 5(5):III, 1994), to Lake Famur from nearby fish farms (Maafi, 1996b), to the Kor River basin in Fars (A. Alamdari, in litt., 1997; M. Hafezieh, in litt., 1997), throughout Khuzestan Province (not caught in the 1970s), to the Kol River, Hormozgan (Bagheri et al., 2010), and to numerous other water bodies and ponds throughout Iran where it reproduces in the wild. Mirror carp found in the Shadegan marshes of Khuzestan are escapees from fish farms. Carp in the Bahu Kalat and Sarbaz rivers of Baluchestan are also escapees from fish farms (A. Mobaraki, pers. comm., June 1999).

Abdoli (2000) records this species as present generally in the Dasht-e Lut, Sistan, Dasht-e Kavir and Kerman-Na'in basins; in lower reaches of rivers from the Gorgan to the Astara and along the Caspian coast, the middle Aras River; the Kashaf River in Khorasan; the middle and lower Talkheh River and Tatavi River in the Orumiyeh basin; the Qareh Chai and Qom rivers in the Namak Lake basin; the middle to lower Zayandeh River in the Esfahan basin; the Aravand, lower Karun, Jarrahi, lower Karkheh rivers in the Tigris River basin; the middle Halil River in the Hamun-e Jaz Murian basin; and the middle and upper Kul River including the Shur River tributary in the Hormuz basin. Jalali et al. (2005) records this species from the Dez, Karun and Zayandeh rivers, the Vahdat reservoir, and the Zarivar and Kaftar lakes. Abdoli and Naderi (2009) record it from the Atrak, Gorgan, Ghararasu, Tajan, Babol, Aras and Safid rivers, the Anzali Talab, Gorgan Bay, and the southwest, southeast and south-central Caspian Sea.

Zoogeography

The natural distribution of the carp is supposed to be Asia Minor and the Caspian Sea basin where its origins lie in the late Pliocene. From this area, modern wild carp spread east and west, perhaps as late as the last postglacial thermal optimum, and latterly aided by man (Balon, 1974; 1995; Van Damme et al., 2007). Iranian carp may be a mixture of native and introduced stocks (or species - see Kottelat (1997) - but this remains unresolved). It is probably not now possible to distinguish the native stocks morphologically because of admixtures of farmed stocks. All domestic forms probably originated from native Danube River stocks (Bănărescu, Barus and Peňáz in Bănărescu and Paepke, 2002). Khalili and Amirkolaie (2010) compared fish electrophoretically, meristically and morphologically from the Anzali lagoon, Qareh Su and Bandar Gaz. Morphometric differences between east and west were attributed to a relatively smaller body size in the east and the influence of domestic stocks. Electrophoretic differences, particularly obvious at Anzali, were attributed to the larger number of fish farms in this latter area.

Habitat

This species favours an abundance of soft vegetation in shallow water, necessary for successful reproduction. Still waters are preferred but they are found in the lower courses of lowland rivers with moderately flowing water, and occasionally in water exceeding 2 m/sec. They can often be seen basking at the surface or feeding on algae and their dorsal fins break the water surface. Large fish often move into shallows in the afternoon and evening. Carp also leap from the water but the reason is unknown. They rarely descend below 30 m in lakes and avoid fast water in streams. Carp overwinter in depressions in the lower reaches of rivers in the Caspian basin. There are both freshwater resident populations in the Caspian basin and diadromous (or semi-diadromous) ones, the latter living in brackish water near river mouths and only entering fresh water to spawn. The migration up the Ural River may reach 60 km from the river mouth. Some fish apparently spawn in the brackish waters of shallow coastal areas in the Caspian. Knipovich (1921) reports fish in the Iranian Caspian Sea down to 11.9-12.3 m. Riazi (1996) and Karimpour (1998) report that this species is both native (resident) to, and migrates to, the Siah-Keshim Protected Region of the Anzali Mordab.

Carp have a salinity tolerance under experimental conditions of up to 8‰, and for short periods 18.6‰ with acclimation, and this has significance for survival of carp in the Caspian Sea and in waters of southern Iran and Iraq where this species is farmed. Hafezamini and Oryan (2002) and Hafezamini et al. (2003) however found that, under experimental conditions, all fish in their study died at 18‰ in less than 12 hours. Carp eggs hatch in water up to 10‰, with the favourable level being up to 6.6‰ (Al-Hamed, 1971).

Low dissolved oxygen concentrations of 3 mg/l are tolerated and levels as low as 0.5 mg/l can be withstood for 2-3 hours. Normal growth has occurred in fish kept at 35°C. The immune response of carp following exposure to the organophosphate malathion has been studied by Soltani et al. (2003).

This species is considered to be a dominant species in the Karun River along with Barbus (= Tor) grypus (Iranian Fisheries Research and Training Organization Newsletter, 17:1, 1997).

Introduced to Iraqi waters in 1960 as juveniles, this species rapidly became established (Ahmed and Taher, 1988). Cyprinus carpio was caught in large numbers in the Shatt al Arab of Iraq down to the estuary after an increase in the discharge of the Tigris River reduced salinity (N. A. Hussain, in litt., 1994).

Age and growth

Fatemi et al. (2009) examined fish from beach seines along the Caspian shore of Iran for 2006-2007. Ten age groups were recorded and the catch was dominated by fish aged 4-5 years old. Growth parameters based on scale reading were FL = 71.52 cm and K = 0.16 per year for the total population, FL = 70.54 cm and K = 0.15 per year for males, and FL = 72.00 cm and K = 0.16 for females. Growth parameters based on length-frequency analysis gave values of FL = 72.0, 69.3 and 73.0 cm and K = 0.18, 0.15 and 0.18 per year respectively. The total (Z), natural (M) and fishing (F) mortalities were 0.71, 0.29 and 0.42 per year respectively for sexes combined. Exploitation (E) was 0.59 for sexes combined and no further fishing pressure was recommended.

Females are larger and mature a year later than males. Sexual maturity is attained in the second year of life, and in a few individuals even by the end of the first year, in the southeastern Caspian Sea; but in the southwestern Caspian Sea this occurs in the third and fourth years (Kuliyev and Agayarova, 1984). Resident carp in Dagestan mature in their third year at about 30 cm and have an average life span of 6 years whereas the semi-anadromous or semi-diadromous form matures in its fourth year at 35-36 cm and has an average life span of 8 years (Shikhshabekov, 1969). Growth is faster in the Kura River of Azerbaijan than in other populations in the Caspian but maturity is later at age 4 or more usually at 5 years. Freshwater residents in the Anzali Mordab are slow-growing compared to the semi-diadromous form and are less common. Fish taken in the commercial operations in Iran were 3-7 years old, 31.0-63.0 cm long and weighed 539-3375 g (Razivi et al., 1972). Mousavi Gel Sefid et al. (2007) give some growth equations for fish from the Anzali Lagoon.

This species was stocked in the Dukan and Derbendikhan dams of Iraq in the 1960s where fish up to 3 years of age were reported by Ciepielewski et al. (2001). Decreasing growth rates indicate conditions are not too favourable although growth in the first two years is comparable with that in lakes of central Iraq. The condition coefficient (K) was higher among smaller fish, e.g. fish from Dukan at 230 g had a K of 2.32, at 1 kg K was 1.75. Ahmed and Taher (1989) examined the growth of 0+ carp in Hawr al Hammar, Iraq found the length-weight relationship to be W = 0.00004627 L2.8022 and derived the growth equation Lt = 189.87 (1-e-0.0158 (t + 25.662)) with a calculated length at the end of the first year of life of 189.87 mm. This is relatively larger than for other parts of the world, indicating a successful introduction of this exotic. Al-Nasiri and Dawood (1991) found the smallest mature male was 182 mm and the smallest female was 184 mm in Hawr al Hammar, with sexual maturity achieved in the first year of life. Maximum life span was 8+ years. Epler et al. (2001) found the oldest age groups in Iraqi lakes to be 5+ in Lake Habbaniyah and 3+ in Lake Razzazah. The mean condition factor was 1.47 for Lake Habbaniyah and 1.50 for Lake Razzazah.

In Sariyar Dam Lake near Ankara in central Anatolia, ages range from 0 to 18 years (Ekmekci, 1996b). In their first year, fish have an average fork length of 103 mm and weigh 24 g, in 5 years they average 357 mm and 822 g, and in 10 years 580 mm and 3365 g. In Gölhisar Lake by contrast, a small water body in western Turkey, age composition was from 1 to 6 years and fish attained a maximum of 494 mm and 1922 g (Alp and Balik, 2000).

Maximum life span for this species is reported as 47 years for domestic fish.

Food

Food is derived from browsing on the substrate at all hours, if the temperature is favourable. Browsing muddies the water and can inhibit other species and uproot plants. Mouthfuls of bottom ooze are taken up, spat out and the food items selected. These include aquatic insects, crustaceans, worms and molluscs, and more rarely, fish. Plant material is ground up by the molar pharyngeal teeth and includes algae, seeds, wild rice, leaves and various aquatic plants. Organic sewage is also eaten. Some surface feeding on algal mats or insects will also occur. Feeding almost completely stops in winter and the fish go into a form of hibernation.

Feeding in the Hawr al Hammar, Iraq is related to temperature, the peak intensities being July and the minimum in January with peak activity in September and minimum in January. Feeding does occur year round and smaller fish (<200 mm) have highest feeding activity in spring while adults have this in summer (Hussein et al., 2000a). In a study of the recovering Hawr al Hammar, diet was 25.45% algae, 18.18% snails, 12.73% diatoms and 12.73% copepods, 10.91% insects with plants, cladocerans and shrimps at less than 10% each, in the Hawr al Hawizah diet was 27.3% snails, 18.2% insects and 12.1% for algae, plants and cladocerans with fish, diatoms and copepods at less than 10% each, and in the Al Kaba'ish (= Chabaish) Marsh 33.3% algae, 20.4% insects, 11.1% snails and diatoms and plants at 10.2, with various crustaceans at less than 10% each (Hussain et al., 2006). Hussein et al. (1991) examined diet in the Garma Marshes, Iraq and found crustaceans, molluscs, aquatic plants and seeds, aquatic insects, oligochaetes and fish to be dietary items, selection and numbers varying with carp size and season. Some fish were found to have fed exclusively on only a single, different mollusc species, presumably as opportunity presented. Al-Shamma'a et al. (1996) examined the food of this species in Al Qadisiyah Reservoir, Iraq and found plants, their seeds, molluscs and aquatic insects, all bottom foods. Mangalo and Akbar (1988a, 1988b) studied the food of carp in a farm pond at Al-Latifiyah, Baghdad where zooplankton was the principal diet. The gill rakers show an efficient structure for filtration, indicative of a phytoplanktivorous and omnivorous feeding (Salman et al., 1993). Salman et al. (1994) report a mixture of animal and plant foods, with zooplankton a dominant component of all length groups. The gut is coiled and 3.42 times standard length, indicating omnivory with plant food being important. Epler et al. (2001) gave the diet in Lake Habbaniyah, Iraq as 51.7% plants, 15.7% oligochaetes, 15.2% tendipedids, 7.2% molluscs, 5.2% detritus and 4.1% cladocerans. They also found that where there is significant competition between autochthonous species, as here, carp become another strong competitor for food. Dietary coincidence between carp and shabbout, gattan and himri was 58.5, 68.5 and 54.2 % respectively. Hussein et al. (2000b; 2000c) examined dietary overlap between this species and three native carps in the Hawr al Hammar. Overlap with Barbus (= Mesopotamichthys) sharpeyi was the weakest as this species is a herbivore but small B. luteus (<200 mm) and B. xanthopterus showed strong overlaps. This overlap may explain the decline in some native carps. Hussain and Ali (2006) also examined feeding relationships among fishes in the Hawr al Hammar and found this species to be a carnivore, 26.4% of the diet being crustaceans, 12.7% insects and 30.5% molluscs. Dietary overlap of 84% was found between this species and Barbus (= Luciobarbus) xanthopterus but the availability of food resources offset possible competition, contrasting with the conclusions above. Ciepielewski et al. (2001) found the diet of this species in the Dukan and Derbendikhan reservoirs in Iraq to be mainly algae, copepods and chironomids.

Reproduction

Under natural conditions, males spend more time on the spawning grounds than females and spawn several times. More than 7 million eggs up to 1.71 mm in diameter may be present in a female but only about 500 are laid at a time. Wild carp in the Atrek River had a fecundity range of 16,000 to 543,000 eggs (Bănărescu, Barus and Peňáz in Bănărescu and Paepke, 2002). The resident form is less fecund by about half than the semi-anadromous form (Shikhshabekov, 1969). In the Anzali Mordab a mass spawning run takes place in April with spawning in April-May. The first migratory fish are seen as early as January. Shallow weedy areas or the mouths of rivers are used as spawning sites and adhesive eggs are laid on plants.

Resident populations in Dagestan spawn earlier, by about a month, than the semi-anadromous population which spawn in early May (Shikhshabekov, 1969). Spawning time variations are governed by temperature and the most favourable temperature is 18-20°C.

Carp ascend the Kura River of Azerbaijan in spring and autumn. The spring run begins in mid-March and peaks in April while the weak autumn run lasts from August to mid-October.

Carp have feeding grounds in the coastal waters of the southeastern Caspian Sea and enter the Atrak River in winter to spawn between February and April. Young migrate downstream, this movement ending in July when the river flow is minimal or ceases. When there is no flood, spawning does not occur (Petr, 1987). Ghelichi et al. (2010) used histology and gonadosomatic indices on fish from the commercial fishery in Golestan and from research organizations to determine that spawning occurred over at least 8 months. The mean absolute fecundity was 143,302 eggs. Shirali et al. (2011) using histology and histometry on broodstock found a breeding season from April to October in Khuzestan. The climate allows rapid maturation and the ability to spawn three times in a year.

Fish in Iraqi ponds grew 25-30 cm in the first year of life and matured in 1-2 years. At 16-26ºC they spawned from late February to late April and again in the autumn (Al-Hamed, 1960). Palm tree fibres were used for egg deposition and eggs hatched in 4-8 days. Al-Nasiri and Dawood (1991) found a fecundity range of 14,150-1,492,500 eggs in Hawr al Hammar with a mean relative fecundity of 182 eggs/g of body weight, and a egg diameters of 0.90-1.02 mm. The gonadosomatic index indicated spawning in March and possibly October-November. Epler et al. (2001) studied reproduction in lakes Tharthar and Habbaniyah and found both sexes to achieve maturity in the first year of life at 13.5 cm for males and 12.6 cm for females. Spawning occurred in May and fecundity was 186-531 thousand eggs/kg body mass.

The spawning behaviour involves stimulation of a female while moving over vegetation and being accompanied by 2-3 males, active movement and spawning being induced by blows from the male(s). The eggs adhere to the vegetation or are lost. Most eggs are shed at night or in the early morning.

Shirali et al. (2012) give a histology of ovarian development in carp from culture ponds in Khuzestan.

Parasites and predators

Eslami and Anwar (1971) record the cestode Caryophyllaeus fimbriceps from this species on the Caspian coast of Iran. Mokhayer (1976b) records the cestodes Bothriocephalus gowkongensis, the nematode larva Anisakis and the acanthocephalan Pomphorhynchus perforator. Mokhayer (1989) reports metacercariae of the eye fluke, Diplostomum spathaceum from this species in Iran, which can cause complete blindness and death in commercially important species. Jalali and Molnár (1990b) variously record the monogeneans Dactylogyrus anchoratus, D. extensus, D. sahuensis and D. vastator from carp on fish farms throughout Iran and Naem (2002) records D. anchoratus from fish in the Safid River.

Moghainemi and Abbasi (1992) record a wide range of parasites from this species in the Hawr al-Azim in Khuzestan. Mortazaei et al. (2000) record an infection rate of 66% (2 of 3 fish) with the worm Bothriocephalus opsariichthydis in Khuzestan marshes. Sattari and Faramarzi (1997) record Caryophyllaeus fimbriceps, C. laticeps and C. brachycollis from 38% of carp in the Anzali lagoon. Akhlagi (1999; 2000) reports that high temperatures (up to 32°C) stresses this species and leaves it open to infection with Aeromonas hydrophila. Safari and Khandagi (1999) record Clostridium botulinum from 3.8% of fresh and smoked samples of this species in Mazandaran Province. Mousavi and Khosravi (1999; www.mondialvet99.com, downloaded 31 May 2000) record the toxigenic fungi Aspergillus flavus, Alternaria, Penicillium and Fusarium from this species and the pond water at a fish farm in northern Iran. Farahnak (2000) records Anisakidae from this species in Khuzestan. Akhondzadeh et al. (2002) and Akhondzadeh Basti and Zahrae Salehi (2003) show that the psychotropic pathogen Listeria monocytogenes is found in market and fish farm samples of this species. Masoumian et al. (2002) investigated parasites from this fish in the Aras and Mahabad dams in northwest Iran and found the protozoan Goussia carpelli, also known from carp in the Safid River. Ebrahimzadeh Mousavi et al. (2005) isolated the fungus Branchiomyces spp. from gill lesions of farmed carp in northern Iran. Branchiomycosis or gill rot is a major problem in commercial fish production.

The intestinal helminth Bothriocephalus gowkongensis was recorded from this species on fish farms in West Azarbayjan Province (Azarvandi et al., 1999). Naem et al. (2002) found the following parasites on the gills of this species from the western branch of the Safid River, namely the protozoans Ichthyophthirius multifilis and a Trichodina species, a copepod crustacean Lernaea sp., monogenean trematodes Dactylogyrus anchoratum, and D. achmerowi. Jalali et al. (2002) and Jalali and Barzegar (2006) record parasites from this species in Lake Zarivar, namely Trichodina pediculus, Dactylogyrus extensus, Gyrodactylus stankovici, Diplostomum spathaceum two species of Argulus, Pseudocapillaria tomentosa and Lernaea cyprinacea. Jalali et al. (2005) summarise the occurrence of Gyrodactylus species in Iran and record G. cyprini, G. elegans, G. shulmani, G. sprostonae, G. stankovici and G. sp. from various localities for the carp. Farahnak et al. (2002) record Contracaecum sp. and Anisakis sp. from this fish in Khuzestan Province. Mehdipoor et al. (2004) record the monogenean Dactylogyrus extensus in Zayandeh River fish. Araghi Soureh and Jalali Jafari (2005) recorded Dactylogyrus extensus from this species in the Mahabad River of the Lake Orumiyeh basin. Jalali and Barzegar (2005c) record five species of monogeneans in the genus Dactylogyrus from both farmed and native Cyprinus carpio in Iran. These are D. extensus, D. anchoratus, D. achmerovi, D. vastator and D. sahuensis. Fry and fingerlings are more sensitive to these parasites and this sensitivity is increased with crowding in ponds. The paper also deals with gill histopathology and distribution of the parasites in Iran. Barzegar and Jalali (2006) report parasites in this species from Kaftar Lake as Lernaea cyprinacea,  Trichodina sp., Dermocystidium sp., Dactylogyrus extensus, D. anchoratus, Gyrodactylus sp. and Diplostomum spathaceum. Khara et al. (2006b) record the cestode Caryophyllaeus fimbriceps from this species in the Boojagh Wetland of the Caspian coast. Pazooki et al. (2007) recorded various parasites from localities in West Azarbayjan Province, namely Diplostomum spathaceum, Ligula intestinalis, Digrama sp. and Argulus foliaceus. Sattari et al. (2007) record the cestode Caryophyllaeus fimbriceps, the digenean Diplostomum spathaceum and the monogeneans Dactylogyrus extensus, Gyrodactylus sp. and Diplozoon sp. in this species in the Anzali wetland of the Caspian shore and also mention that the Caryophyllaeus laticeps is also known from this species in the Iranian Caspian Sea. Barzegar et al. (2008) record eye parasites from this fish including the monogenean Gyrodactylus stankovici, the digeneans Diplostomum spathaceum and Tylodelphys clavata, and the crustacean Lernaea cyprinacea. Khara et al. (2008) found the eye parasite Diplostomum spathaceum in this fish from Boojagh Kiashar Wetland in Gilan. Shamsi et al. (2009) found Dactylogyrus achmerovi, D. anchoratus, D. extensus, D. sahuensis and D. vastator in this species in fish farms, Safid River and Hamun Lake. Barzegar and Jalali (2009) reviewed crustacean parasites in Iran and found Argulus sp., Argulus foliaceus, Ergasilus sieboldi, Lernaea sp. and Lernaea cyprinacea on this species. Khara et al. (2011) list the mongenean Dactylogyrus sp., the digenean Diplostomum spathaceum, the cestode Caryophyllaeus fimbriceps, the crustacean Lernaea cyprinacea and the leech Piscicola sp. from this fish in the Boojagh Wetland of the Caspian Sea. Maktabi et al. (2011) recorded the incidence of Listeria spp. on Khuzestan market fish from fish farms and the frequency was a matter of concern..Abedi et al. (2012) reported the first epidermal inclusion cyst in a pond specimen of carp.

The Caspian seal, Pusa caspica, is a significant predator on this species (Krylov, 1984). Esox lucius, Sander lucioperca and Silurus glanis are all predators on small carp. A wide variety of other fishes and birds eat smaller carp as do predatory aquatic insects, frogs and toads. Carp eggs are eaten by many fishes. Ashoori et al. (2012) found that grey herons (Ardea cinerea) in the Siahkeshim Protected Area of the Anzali Wetland ate this species. Adult carp, however, are too large for most predators to take.

Economic importance

Balon (1974; 2006) details domestication of the carp which has been a cultivated fish for over 2000 years. The carp is an important table fish in Iraq where details of farming techniques are given by Ahmed and Taher (1988) and by Mangalo and Akbar (1988a; 1988b). Catches in former Soviet waters of the Caspian Sea were described by Kuliyev and Agayarova (1984). In the southwestern part, size range was 21-64 cm in 1960-1961 (92% of fish were 29-41 cm) but in 1972 95% of fish were 31-55 cm. The smallest carp are caught in the southeastern Caspian where maximum weight was 3.97 kg as opposed to 10.0 kg in the central Caspian. Spawning fish in the southeastern Caspian were only 0.58-1.0 kg while in the southwestern Caspian carp were 0.25-5.75 kg. About 17-30% of commercial catches in the Volga region are estimated to be comprised of hatchery production (Petr, 1987).

Nevraev (1929) gives catches for various fishing regions in Iran in the early twentieth century. For the Safid River region from 1899-1900 to 1917-1918 the catch was 30 to 30,500 individuals with no fish reported in some years, and in Astrabad (= Gorgan) region from 1900-1901 to 1912-1913 the catch was 14,200 to 851,800 individuals. The commercial catch in Iran from 1956/1957 to 1961/1962 varied from 3443 kg to 175,295 kg (Vladykov, 1964), from 1965/66 to 1968/69 varied from 184 to 333 tonnes (Andersskog, 1970) and from 1963 to 1967 from 7.0 to 108.8 t (on a yearly basis 41.1, 9.8, 7.0, 108.8, and 48.1 t respectively) (RaLonde and Walczak, 1970b). Catches from 1933/34 to 1961/62 varied from 34 kg to 1113 t in the Bandar Anzali region and the total catch of the Northern Shilot (Fisheries Company) varied from 9.8 t to 333 t from 1963/64 to 1968/69 (RaLonde and Walczak, 1972). The Food and Agriculture Organization, Rome gave the catches from Iran for the 6 years 1980 to 1985 as 1032, 2000, 1000, 52, 83, and 100 t respectively. The catch in the Anzali Mordab in 1990 was 6855 kg and from 1932 to 1964 the annual catch varied from 1.2-638.6 t (Holčík and Oláh, 1992). These wide fluctuations in reporting and in nominal catches are indicative of the poor fisheries statistics as well as the inter-year variability of catches in Iran.

March and April are the main fishing months in Iran (Farid-Pak, no date). It is the main fish in Iranian carp farms, when raised in polyculture with Chinese carps (Jalali and Molnár, 1990b). Salehi (2004) gives a figure of 25% as the share of common carp in Iranian polyculture (63% being silver carp) and gives a review of the economics of carp polyculture. Kohnehchahry and Heydarpur (1973) outline methods of raising carp using submerged cages which they believe would be suitable for Iranian waters. There were about 4000 ha of carp fish ponds (which presumably included Chinese carps such as Hypophthalmichthys spp.) with an annual production of 2-3 tonnes/ha (White, 1988). Plans were made in the 1980s to increase total pond area (including trout which was at 60 ha and produced 1000 t/year) to 35,000 ha over 10 years to yield an annual harvest of 100,000 t/year. However Bartley and Rana (1998b) report an aquaculture production of 6561 tonnes in 1995 and Salehi (2004) gives 28,060 t from warm-water fish farming, mostly Chinese carps. More than 700,000 carp fingerlings were released in the small and remote province of Chahar Mahall va Bakhtiari alone and 20 million carp, silver carp and grass carp fingerlings were produced in the Shahid Rajaae Hatchery in Sari for release across Iran in reservoirs and dams (Abzeeyan, Tehran, 4(7):VII, 1993). Salehi (1999) states that the marketable size of cultured carp is 1 kg with most harvested once annually and almost 90% supplied to market in October-March with a peak in March (the Iranian New Year when fish is a traditional food).

Hasani et al. (2012) investigated the quality of fish fingers produced from carp fillets and surimi and stored in a refrigerator, determining a shelf life of 6 days.

Marjan Iran Company was selling 1500-1800 g fish for U.S.$1.90/kg in August 2003 compared to $2.10 for Ctenopharyngodon idella (http://groups.yahoo.com/groups/hilsa/message/25). In Golestan Province, carp cost 1500-1700 tomans/kg in the early 2000s. Salehi (2006a) analysed the consumer market for this species and its products in Iran. Shabanpour et al. (2007) investigated preparation of surimi from this species.

An exotic species in some parts of the world, carp are a nuisance because they uproot vegetation used by native species for cover, food and spawning. This activity also increases water turbidity to levels which many native species cannot tolerate. Stirred up silt may smother eggs of native species. Carp also compete with other species for food and eat the eggs of other species.

Various studies have been carried out on its culture in Iran and neighbouring countries, e.g. see Freshwater Fishes of Iraq, and its use as an experimental organism, e.g. in Iran, see Mohagheghi and Hedayatifard (Farsi translation of Horváth et al. (1985a and 1985b); Khazraiinia et al. (2000) on acute ammonia toxicity; Peyghan et al. (2001) on use of ketamine and xylazine hydrochloride as anaesthetics; Rostami Bashman et al (2001) on histopathological lesions after exposure to copper, zinc, mercury and cadmium compounds; Soltani et al. (2001) on the anti-fungal effectiveness of formalin on hatch rate of eggs; Abtahi et al. (2002) on clove oil having no significant difference with MS222, another anaesthetic used in fish farms; Arabi and Heydarnejad (2002) on the deleterious effects of copper and mercury, used in combating algal blooms and weeds, on the gills of carp; Baradaran Noveyri et al. (2002) on the effects of cryopreservation on motility of spermatozoa; Peyghan et al. (2002) on differences between gonads of fishes from two culture seasons; Dorafshan et al. (2003) on induction of spawning using pituitary extract and GnRH analogue in combination with domperidone; Sharifpour et al. (2003) on the anaesthetic effects of clove oil under various pH and temperature regimes; Vahabzadeh Roodsari et al. (2003) comparing malachite green and hydrogen peroxide to control fungal infections (the latter is effective and less dangerous); Akhlaghi et al. (2004) on phagocytosis in relation to immunostimulants; Ebrahimi (2004) on the deleterious effects of copper, a pollutant, on sperm anatomy; Moini and Basimy (2004) on production of fish cake according to various recipes and its shelf life; Sharifpour (2004) on the histology of the response and the circumstances of wound healing; Soltani et al. (2004) on the effects of clove oil anaesthesia on haematological parameters, serum enzymes and some tissues, up to 200 p.p.m. being deemed safe; Oryan et al. (2005) on  the effects of baclofen on the pituitary system; Sharifpour et al. (2005) on the highly toxic effects of the pesticide endosulfan (with an LC50 of less than 0.1 mg/l); Yousefian (2005) on generating gynogenetic carp through irradiation; Baradaran Noveiri et al. (2006) on cryopreservation of spermatozoa using different extenders; Ghiasi and Mirzagar (2006) on lysozyme content in sub-lethal concentrations of cadmium; Kazemipour and Keivany (2005) on the use of garlic, mallow and motherwort in healing superficial wounds - garlic reduced recovery by one week; Kazemipour et al. (2006) on carp kept in aquaria at low concentrations of garlic (0.1 g/L) which healed superficial wounds more quickly than controls or aquaria having mallow or motherwort; Kohodabandeh and Abtahi (2006) on the use of sodium chloride, iodine and formalin to control Saprolegnia sp. on eggs (sodium chloride was recommended); Naji et al. (2007) on the 96h LC50 value of cobalt chloride at 327-328 mg/l; Naji et al. (2007) on the toxic effect of zinc sulphate on gill tissues which suffer hypertrophy and hyperplasia; Darafsh et al. (2008) on the use of scales as an indicator of heavy metal pollution; Golchinrad et al. (2008) on the effects of detergent on liver glycogen (decreased) and glucose (increased); Hasanabadizadeh et al. (2008) on the lack of improvement in wound healing after vitamin injections; Imnapour and Enayat Gholampour (2008) on the effect of broodstock migration time on various egg characters; Safari et al. (2008) on changes in muscle chemistry during maturation; Soleymani et al. (2008) on the effectiveness of vitamin C injections on survival of juveniles challenged by different doses of the theront of Ichthyophthirius multifilis; Darvish Bastami and Imanpour (2009) on sperm motility influenced by high concentration of ions; Baghfalki et al. (2009) on the effect of broodstock density on survival and growth of larvae and fingerlings in earthen ponds; Erfani Majd et al. (2009) on evaluating the response of incubated ovarian follicles to carp pituitary extract and cultivated pituitary cells secretion; Ghanbari et al. (2009) on the long term effects of changes in pH on haematological parameters in fingerlings; Imanpour and Safari (2009) on the effect of maturation stages on gonadal indices and chemical composition of the gonad; Imanpour et al. (2009) on stocking density and its effect on survival and growth in polyculture, up to 450 fish per hectare being optimal; Imanpour et al. (2009) on the effects of broodstock age on various egg dimensions, females age 3-6 years being found suitable for propagation; Kordjazi et al. (2009) on physicochemical parameters of water and their correlation with haematocrit indicators, growth and survival in farm ponds; Rostami Mina and Soltani (2009) on the histopathological effects of the aquaculture pollutant copper sulphate; Salamat et al. (2009) on pituitary primary cell culture and its secretion effect on endocrine activity of incubated ovarian follicles; Sheykhzadeh et al. (2009) on the effects of Eucalyptus essential oil on immunological variables; Yehganeh et al. (2009) on seasonal variation in the chemical composition and fatty acid profile of ovaries, necessary for embryogenesis; Alishahi et al. (2010) on feeding Aloe vera to enhance immune responses; Baghfalaki et al. (2010) on on food habits of larvae and fingerlings in earthen ponds; Ghiasi et al. (2010) on the effects of low concentrations of cadmium on the immune response in winter; Kordjazi and Imanpoor (2010) on pH and ions in the water and their effects on blood serum in pond fish; Nematollahi (2010) on stress responses to confinement being similar to other fish species; Rafiee and Hekmat (2010) on on the usefulness of populus shavings as a biofilter medium in a recirculating culture system; Rahmanifarah et al. (2010 on stunning and killing fish with clove oil preserves the flesh quality better than other methods such as  carbon dioxide bath and asphyxia; Salamat et al. (2010) on ovarian follicular cells and their endocrine activity in cell culture; Soltani et al. (2010) on the immune responses to Zataria multiflora (Persian thyme) essential oil used as an anti-fungal for carp eggs; Yeganeh et al. (2010) on pre-spawned carp showing a higher variation in lipid quality as frozen fillets; Zolfaghari et al. (2010) showing that fish size has a positive relationship with lipid content and fillet dry weight yield; Afkhami et al. (2011) on a comparison of chemical composition with Ctenopharyngodon idella, there being significant differences in protein, lipid and moisture but not in ash; Banaee et al. (2011) on haematological and histopathological effects of diazinon pesticide; Faramarzi et al. (2011) on the reduction in cost of fish feed with sweet potato peels incorporated; Farhoudi et al. (2011, 2011) on changes in the fatty acid profile of larvae during development in order to determine nutritional requirements and improve product quality; Ghovati et al. (2011) on changes in hardness and alkalinity on toxicity of zinc; Iranshahi et al. (2011) on the immuno-stimulatory effects of prebiotic bacteria and vitamin C on fingerlings; Moini et al. (2011) on stress responses and meat quality after fish were stunned by ice asphyxia, air asphyxia and immersion in clove oil, the latter being best; Pazooki et al. (2011) on heavy metal (lead, zinc and copper) levels in cultured carp not being a health problem; Peyghan et al. (2011) on the detection of the illegal antibiotic furazolidone in cultured carp in Khuzetsan; Salati et al. (2010) on changes in blood parameters used to monitor salinity effects; Seifi et al. (2011) on the use of ovaprim and pituitray extract increased sperm quality over the use of HCG hormone; Shabanpour et al. (2011) on overcrowding and asphyxia cause maximum stress and reduce flesh quality; Sheikhzadeh et al. (2011) on the effects of essential oils from Zataria multiflora and Eucalyptus globolus on haematological parameters and respiratory burst activity in relation to health during temperature stress; Shirali et al. (2011) on on histological development of the ovary during the breeding season; Yousefian et al. (2011) on heritability of growth-related traits in wild carp; Zabihi et al. (2011) on the useful effect of sodium selenite as a diet additive for carp fingerlings; Banaee et al. (2012) on on the effects of the agricultural pesticide diazinon on biochemical blood parameters and liver histopathology; Dehghani et al. (2012) on the copper bioaccumulation patterns of common and mirror carp; Khajepour et al. (2012) on a comparison of juvenile diets involving crude protein, citric acid and microbial phytase; Kordjazi et al. (2012) on a salinity of 0.3-2.7 g/l and an electrical conductivity of 843-5230 μm/cm2 without stress favoured growth; Peyghan et al. (2012) on diet cholesterol used to increase the amount of sex hormones; Salamat et al. (2012) on synthetic carp GnRH and carp pituitary homogenate for in vivo induction of ovulation and spawning; Sanchooli et al. (2012) on on the role of epidermal mucus and its components in the immune system; Shamoushaki et al. (2012) on management and optimisation of feeding frequency for juveniles; etc.

This species is actively angled for along the Caspian shore of Iran and in its rivers (e.g. see Noorbakhsh (1993a)), appears regularly in fish markets of Ahvaz, Khuzestan and is caught by anglers there using bread or potato as bait.

It is characterised as a fatty fish according to a lipid content 9-14% by wet weight of muscle in autumn (Hantoush et al., 1999). Al-Aswad et al. (1980) detailed the chemical composition of this species in Dukan Lake, Iraq including seasonal levels of moisture, fat, protein and ash, and the various types of fatty acids and amino acids. Hindi et al. (1996a) give the chemical composition of flesh of this species as 78.87% moisture, 2.46% fat, 17.06% protein and 1.35% ash, indicating a valuable food fish characterised as lean to medium fatty. Hindi et al. (1996b) give chemical indices for assessing fish freshness according to the month of capture and marketing (pH 6.28, total volatile nitrogen bases 11.07 mg/N/100g fish, thiobarbituric acid 0.47 mg, and free fatty acids 0.62%). Elsagh (2011) found Iranian fish fillets with cadmium and lead levels above accepted limits for human consumption;

The roe or eggs of this species have been implicated in poisoning (Halstead, 1967-1970; Coad, 1979b) and should be avoided (see under the genus Schizothorax for more information on egg poisoning). Fish should be carefully cleaned in the spawning season to remove the eggs and ensure against contamination of flesh. Severe cases of egg poisoning in other species have resulted in death. Vaezzadeh et al. (2008) found the levels of the pesticide heptachlor in fish from Anzali and Ramsar could have a health risk to consumers.

Robins et al. (1991) list this species as important to North Americans. Importance is based on its use in aquaria and aquaculture, as food, in textbooks, for sport, as an experimental species and because it has been introduced outside its natural range. Brightly coloured varieties of carp are known as "koi" and are kept as ornamental fish. Colours include red, orange, white, black, blue and yellow in various combinations.

Conservation

Vladykov (1964) recommended that fishing for this species in Mazandaran and Gorgan be prohibited for 5 years because of reduced stocks. Krasznai (1987) and Petr (1987) give details of fish farms propagating this species in Iran. For example, 30 million fish were produced by the Safid Rud Fish Farm in 1986. In 1999-2000, 20 million juveniles were released into the Caspian Sea (Iranian Fisheries Research Organization Newsletter, 23:4, 2000). From October to March 2000, 3 million juveniles raised in the Shahid Ansari aquaculture and breeding centre in Gilan were released into the Caspian Sea and neighbouring water bodies (Iranian Fisheries Research Organization Newsletter, 26:2, 2001). Poaching was a problem in the Caspian basin of Iran (Razivi et al., 1972) and no doubt continues. The Atrak River stocks are an important fishery for both Iran and Turkmenistan but are susceptible to loss through the absence of flooding of the spawning grounds. Fish passes are needed to ensure access to the spawning grounds, timely release of water from a reservoir to flood the spawning areas in years of low water flow, enforcement of catch limits, and continued stocking (Petr, 1987). Yousefian et al. (2009) examined the normative reproduction values and genetic characters for native carp in the Caspian Sea. The average weight of sampled fish was 1441.6 g and standard length was 50.5 cm. Fertilisation rate was 62-79%, absolute fecundity was 114,347 and relative fecundity 82,007. There was no significant differences in haplotype distribution between areas in the southern Caspian Sea.

The study of Ghelichi et al. (2011) mentioned above, showing different populations, has implications for conservation and management. Yousefian (2011a) found low heterozygosity using 5 microsatellite loci, perhaps indicative of overfishing, such information being important in managing the stocks.

Lelek (1987) considered that the wild form, as opposed to domestic stocks, was vulnerable to endangered in Europe because of habitat modifications. 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, sport fishing, abundant in numbers, habitat destruction, widespread range (75% of water bodies), present in other water bodies in Iran, and present outside the Caspian Sea basin. Karami et al. (2005) found river fish had greater exposure to organophosphorus compounds than Caspian Sea and well-fed fish.

Further work

Where this species is introduced, its interactions with native species should be carefully studied and monitored in an effort to protect the native fauna.

Sources

A recent summary on this species is Bănărescu, Barus and Peňáz in Bănărescu and Paepke (2002).

Iranian material: CMNFI 1970-0510, 1, 47.8 mm standard length, Gilan, Golshan River (37º26'N, 49º40'E); CMNFI 1970-0522, 1, 108.5 mm standard length, Gilan, Safid River at Astaneh Bridge (37º16'30"N, 49º56'E); CMNFI 1970-0563, 3, 28.6-58.6 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0568, 1, 125.8 mm standard length, Gilan, Caspian Sea at Kazian Beach (ca. 37º29'N, ca. 49º29'E); CMNFI 1970-0582, 3, 63.3- 134.0 mm standard length, Mazandaran, Aliabad Reservoir (36º56'N, 54º50'E); CMNFI 1970-0587, 2, 47.5-76.7 mm standard length, Mazandaran, Babol River at Babol Sar (36º43'N, 52º39'E); CMNFI 1979-0455, 6, 46.9-67.1 mm standard length, Markazi, Manjil Dam (36º45'N, 49º17'E); CMNFI 1979-0476, 3, 44.1-85.3 mm standard length, Mazandaran, Qareh Su near Kord Kuy (36º51'N, 54º05'E); CMNFI 1979-0479, 19, 28.0-143.8 mm standard length, Mazandaran, dam on Gorgan River (37º09'30"N, 54º41'30"E); CMNFI 1979-0685, 2, 31.5-45.7 mm standard length, Gilan, Safid River (ca. 37º22'N, ca. 49º57'E); CMNFI 1979-0788, 2, 104.2-123.5 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0128, 3, 60.2-75.2 mm standard length, Mazandaran, Qareh Su (36º49'30"N, 54º03'30"E); CMNFI 1980-0132, 4, 56.0-64.6 mm standard length, Gilan, Safid River at Kisom (37º12'N, 49º54'E); CMNFI 1980-0157, 1, 126.3 mm standard length, Mazandaran, Gorgan River estuary (36º59'N, 53º59'30"E); CMNFI 1980-0905, 2, 78.1-92.9 mm standard length, Mazandaran, Gorgan River at Khadje Nafas (37º00'N, 54º07'E); CMNFI 1980-0908, 1, 54.4 mm standard length, Gilan, Safid River estuary (ca. 37º28'N, ca. 49º54'E).

© Brian W. Coad (www.briancoad.com)