Reproductive biology of the invasive sharpbelly, Hemiculter leucisculus (Basilewsky, 1855), from the southern Caspian Sea basin



The sharpbelly, Hemiculter leucisculus, an invasive species, has expanded its range throughout much of Asia and into the Middle East. However, little is known of its reproductive information regarding spawning pattern and season that could possibly explain its success as an invasive species. This research is the first presentation of its reproductive characteristics, which was conducted based on 235 individuals collected monthly throughout a year from Sefid River, in the southern Caspian Sea basin. Age, sex ratio, fecundity, oocytes diameter and gonado-somatic index were calculated. Regression analyses were used to find relations among fecundity and fish size, gonad weight (Wg) and age. The mature males and females were longer than 93.0 and 99.7mm in total length, respectively (+1 in age). The average egg diameter ranged from 0.4mm (April) to 1.1mm (August). Spawning took place in August, when the water temperature was 23 to 26°C. Average gonado-somatic index values for females at the beginning of the reproduction period (March) and in ripe mature spawning females (August) were 2.90% and 16.97%, respectively. The averages of absolute and relative fecundities were 19978.6 and 607.1 eggs, respectively. The absolute fecundity was significantly related to body weight, ovary weight and age. A potentially high reproductive capacity, comparatively small eggs, short time of eggs’ development, fast sexual maturation, short life span, tolerance to a wide range of environmental conditions, all promote the invasion of this invasive sharpbelly in freshwater ecosystems of the temperate climate zone.


Gonado-somatic index, Fecundity, Freshwater ecosystem, Spawning season.

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Abdoli, A. 2000. The Inland Water Fishes of Iran. Iranian Museum of Nature and Wildlife, Tehran. 378 pp. [In Persian]

Ahmed, A.; Sharaf, M.M. & Laban, H.A 2010. Reproduction of the Egyptian Sole, Solea aegyptiaca (Actinopterygii: Pleuronectiformes: Soleidae), from Port Said, Egypt, Mediterranean Sea. Acta Ichthyologica et Piscatoria 40(2): 161-166.

Ayoade, A.A. 2011. Population characteristics of Schilbe mystus (Linne, 1758), from two different habitats: Asejire and Oyan Lakes Southwestern Nigeria. Journal of Fisheries and Aquatic Science 6(5): 571-577.

Bagenal, T.B. & Braum, E. 1978. Eggs and early life history. In: T. Bagenal (ed.) Methods for assessment of fish production in freshwaters. 3rd edn. Blackwell Scientific, Oxford, UK. pp: 106.

Bagenal, T.B. 1967. A short review of fish fecundity. In: S.D. Gerking (Ed.), The Biological basis of fresh water fish production. Blackwell Scientific, Oxford, UK. pp: 89-111.

Biswas, S.P. 1993. Manual of methods in fish biology. South Asian publishery. New Delhi, pp: 79–91.

Borisova, A.T. 1972. Accidental invaders in water bodies of Uzbekistan. Journal of Ichthyology 12: 49-53.

Cao, Y.P. & Wen, Z.G. 1996. The biology of Hemiculter leucisculus in Bai Yangdian Lake. Journal of Hebei University(Natural Science Edition)m16: 2730. (In Chinese, English abstract)

Chakraborty, B.K.; Mirza, Z.A.; Miah, M.I.; Habib, M.A.B. & Chakraborty, A. 2007. Reproductive cycle of the endangered sarpunti, Puntius sarana (Hamilton, 1822) in Bangladesh. Asian Fisheries Society 20: 145-164.

Coad, B.W. & Hussain, N.A. 2007. First record of the exotic species Hemiculter leucisculus (Actinopterygii: Cyprinidae) in Iraq. Zoology in the Middle East 40: 107-109.

Coad, B.W. 1980. Environmental change and its impact on the freshwater fishes of Iran. Biological Conservation 19(1): 51-80.

Coad, B.W. 2017. Freshwater Fishes of Iran. Available at (accessed on 9 August 2017).

Coad, W.B. & Abdoli, A. 1993. Exotic Fish Species in the Freshwaters of Iran. Zoology in the Middle East 9: 65-80.

Coates, D. 1988. Length-dependent changes in egg size, and fecundity in females, and brooded embryo size in males of fork tailed catfishes (Pisces: Ariidae) from the Sepik River, Papua, New Guinea, with some implications for stock assessments. Journal of Fish Biology 33(3): 455-464.

Cortes, E. 2000. Life history patterns and correlations in sharks. Reviews in Fisheries Science 8(4): 299–344.

Eagderi, S.; Mojazi Amiri, B. & Adriaens, D. 2013. Description of the ovarian follicle maturation of the migratory adult female bulatmai barbel (Luciobarbus capito, Güldenstädt 1772) in captivity. Iranian Journal of Fisheries Sciences 12(3): 550-560.

Esmaeili, H.R.; Gholamifar, A. & Freyhof, J. 2011. Ichthyofauna of Zarivar Lake (Iran) with the first records of Hemiculter leucisculus and Alburnus hohenackeri in the Tigris drainage. Electronic Journal of Ichthyology 7(1): 1-6.

Esmaeili H.R.; Sayyadzadeh G.; Eagderi S. & Abbasi, K. 2018. Checklist of freshwater fishes of Iran. FishTaxa 3(3): 1-95.

Faghani-Langroudi, H. & Mousavi-Sabet, H. 2018. Reproductive biology of lotak, Cyprinion macrostomum Heckel, 1843 (Pisces: Cyprinidae), from the Tigris River drainage. Iranian Journal of Fisheries Sciences 17(2): 288-299.

Fleming, I.A. & Gross, M.R. 1990. Latitudinal clines: a trade-off between egg number and size in Pacific salmon. Ecology 71(1): 1-11.

Gao, X.; Zeng, Y.; Wang, J.W. & Liu, H.Z. 2010. Immediate impacts of the second impoundment on fish communities in the Three Gorges Reservoir. Environmental Biology of Fishes 87: 163-173.

Heidari, A.; Khoshkholgh, M. & Mousavi-Sabet, H. 2014b. Tracing the effects of Sefidrud dams on Capoeta gracilis (Cyprinidae) populations using Truss distances in southern Caspian Sea basin. Iranian Journal of Ichthyology 1(2): 106-113.

Heidari, A.; Mousavi-Sabet, H.; Khoshkholgh, M. & Esmaeili. H.R. 2014a. Morphometric and Meristic Comparison of Capoeta capoeta (Güldenstaedt, 1773) from Upstream and Downstream of the Manjil Dam and Downstream of Tarik Dam in Sefidroud River. Journal of Fisheries 67(2): 207-222. (In Persian)

Heidari, A.; Mousavi-Sabet, H.; Khoshkholgh, M.; Esmaeili, H.R. & Eagderi, S. 2013. The impact of Manjil and Tarik dams (Sefidroud River, southern Caspian Sea basin) on morphological traits of Siah Mahi Capoeta gracilis (Pisces: Cyprinidae). International Journal of Aquatic Biology 1(4): 195-201.

Holcik, J. & Razavi, B.A. 1992. On some new or little-known freshwater fishes from the Iranian coast of Caspian Sea. Folia Zoologica 41 (3): 271-280.

Jamali, H.; Eagderi, S.; Esmaeilzadegan, E. & Patimar, R. 2015. Age, growth and some biological characteristics of Silver bream (Blicca bjoerkna L., 1758) (Cyprinidae) from Aras Dam Lake in Northwest of Iran. International Journal of Aquatic Biology 3(1): 35-41.

Jonsson, N. & Jonsson, B. 1999. Trade-off between egg mass and egg number in brown trout. Journal of Fish Biology 55(4): 767-783.

Jouladeh-Roudbar, A.; Vatandoust, S.; Eagderi, S.; Jafari-Kenari, S. & Mousavi-Sabet, H. 2015 Freshwater fishes of Iran; an updated checklist. AACL Bioflux 8(6): 855-909.

Kolpakov, N.; Barabanshchikov, E. & Chepurnoi, A.Y. 2010. Species composition, distribution, and biological conditions of nonindigenous fishes in the estuary of the Razdol’naya River (Peter the Great Bay, Sea of Japan). Russian Journal of Biological Invasions 1: 87–94.

Li, Q.; Li, W.J. & Zhao, J. 2009. Histological studies on the gonadial development of Hemiculter leucisculus. Journal of Guangzhou University 8: 38-41.

Lobon-Cervia, J.; Utrilla, C.G.; Rincon, P.A. & Amezcua, F. 1997. Environmentally induced spatio-temporal variations in the fecundity of brown trout Salmo trutta l.; tradeoffs between egg size and number. Freshwater Biology 38: 277-288.

Mansouri-Chorehi, M.; Mousavi-Sabet, H.; Sattari, M.; Nasrollahzadeh, A. & Hedayati, A. 2016. Age, sex ratio, spawning season and fecundity of Alburnoides samiii (Pisces: Cyprinidae), from Sefidroud River (the southwestern Caspian Sea basin). Biharean Biologist 10(2): 93-97.

Mohammadi-Darestani, M.; Mousavi-Sabet, H.; Vatandoust, S. & Ahmadnejad, M. 2016. Age, Growth and Reproduction of the Iranian Goby, Ponticola iranicus (Perciformes: Gobiidae) from the Southern Caspian Sea Basin. Journal of Ichthyology 56(4): 578-587.

Moradinasab, Gh.; Daliri, M.; Ghorbani, R.; Paighambari, S.Y. & Davoodi, R. 2012. Length-weight and length-length relationships, Relative condition factor and Fulton’s condition factor of Five Cyprinid species in Anzali wetland, southwest of the Caspian Sea. Caspian Journal of Environmental Science 10(1): 25-31.

Morita, K. & Takashima, Y. 1998. Effects of female size on fecundity and egg size in white-spotted charr: Comparison between sea-run and resident forms. Journal of Fish Biology 35 (5): 1140-1142.

Mousavi-Sabet, H. 2012. Biology of the southern Caspian Spined Loach and it’s keeping in aquarium. Aquariumwereld 65(9): 234-241. (In Dutch)

Mousavi-Sabet, H.; Abdollahpour, S.; Vatandoust, S.; Faghani-Langroudi, H.; Salehi-Farsani, A.; Salehi, M.; Khiabani, A.; Habibi, A. & Heidari, A. 2017. Reproductive biology of Alburnus mossulensis (Teleostei: Cyprinidae) in the Gamasiab River, western Iran. Iranian Journal of Ichthyology 4(2): 171-180.

Mousavi-Sabet, H.; Habibi, A. & Bagherpur, O. 2013. Studies on Length-weight and Length-length Relationships, Relative Condition Factor and Fulton’s Condition Factor of Hemiculter leucisculus (Pisces: Cyprinidae) from the Southwestern Caspian Sea Basin. Our Nature 11 (1): 25–30.

Mousavi-Sabet, H.; Heidari, A. & Paknejad, S. 2015a. Length-weight and length-length relationships of the genus Alosa (Clupeoidei: Clupeiformes: Clupeidae) along the southern Caspian Sea coast. Journal of Applied Ichthyology 32: 129-130.

Mousavi-Sabet, H.; Kamali, A.; Soltani, M.; Bani, A. & Rostami, H. 2012. Age, sex ratio, spawning season, gonadosomatic index, and fecundity of Cobitis faridpaki (Actinopterygii, Cobitidae) from the Siahrud River in the southeastern Caspian Sea basin. Caspian Journal of Environmental Science 10 (1): 15–23.

Mousavi-Sabet, H.; Kamali, A.; Soltani, M.; Bani, A.; Esmaeili, H.R.; Rostami, H.; Vatandoust, S. & Moradkhani, Z. 2011. Age, reproduction, and fecundity of a population of Cobitis sp. (actinopterygii: cypriniformes: cobitidae) from the babolrud River in the southern Caspian Sea basin. Acta Ichthyologica Et Piscatoria 41(2): 117-122.

Mousavi-Sabet, H.; Kamali, A.; Soltani, M.; Bani, A.; Esmaeili, H.R.; Khoshbavar Rostami, H.; Vatandoust, S. & Moradkhani, Z. 2012c. Reproductive biology of Cobitis keyvani (Cobitidae) from the Talar River in South of the Caspian Sea Basin. Iranian Journal of Fisheries Science 11: 383-393.

Mousavi-Sabet, H.; Homayouni, H.; Marjani, M. & Heidari, A. 2015b. Length-weight relationships for 5 Clupeiformes species from the Persian Gulf and Oman Sea. Journal of Applied Ichthyology 32: 169-170.

Nikolsky, G.V. 1963. The ecology of fishes. Academic Press, London.

Patimar, R.; Abdoli, A. & Kiabi, B.H. 2008. Biological characteristics of the introduced sawbelly, Hemiculternleucisculus (Basilewski, 1855), in three wetlands of northern Iran: Alma-Gol, Adji-Gol and Ala-Gol. Journal of Applied Ichthyology 24 (5): 617-620.

Pawson, M.G.; Pickett, G.D. & Witthames, P.R. 2000. The influence of temperature on the onset of first maturity in sea bass. Journal of Fish Biology 56(2): 319-327.

Petr, T. & Mitrofanov, V.P. 1998. The impact on fish stocks of river regulation in Central Asia and Kazakhstan. The Lake and Reservoir Management Journal 3: 143-164.

Pourshabanan, A.; Sari, A. & Mousavi-Sabet, H. 2017. Reproductive biology of female Oxynoemacheilus bergianus (Actinopterygii: Cypriniformes: Nemacheilidae) from the Jajrud River, the Namak Lake basin, central Iran. Iranian Journal of Ichthyology 4(3): 281-289.

Radkhah, A.; Eagderi, S. & Mousavi-Sabet, H. 2016. First record of the exotic species Hemiculter leucisculus (Pisces: Cyprinidae) in southern Iran. Limnetica 35 (1): 175-178.

Rodríguez, L.; Zanuy, S. & Carrillo, M. 2001. Influence of daylength on the age at first maturity and somatic growth in male sea bass (Dicentrarchus labrax, L.). Aquaculture 196 (1-2): 159–175.

Sal’nikov, V.B. 1998. Anthropogenic migration of fish in Turkmenistan. Journal of Ichthyology 38: 591–602.

Tan, X.; Li, X.; Lek, S.; Li, Y.F.; Wang, C.; Li, J. & Luo, J.R. 2010. Annual dynamics of the abundance of fish larvae and its relationship with hydrological variation in the Pearl River. Environmental Biology of Fishes 88: 217–225.

Thrope, J.E.; Miles, M.S. & Keay, D.S. 1984. Development rate, fecundity and egg size in Atlantic salmon, Salmo salar. Aquaculture 43: 289–305.

Thulasitha, W.S. & Sivashanthini, K. 2013. Reproductive characteristics of Doublespotted queenfish, Scomberoides lysan (Actinopterygii: Perciformes: Carangidae), from Sri Lankan waters: implications for fisheries management. Acta Ichthyologica et Piscatoria 43(1): 7–13.

Tourchi-Moghaddam, M. 2003. An investigation on feeding habitats of H. leucisculus in the Anzali lagoon. MSc thesis, Higher Education Institute of Mirza-Kouchakhan, Rasht. pp: 1-137.

Witthames, P.R.; Greer, M.; Walter, M.; Dinis, T. & Whitting, C.L. 1995. The Geographical variation in the potential annual fecundity of Dover Sole, Solea solea, from European Shelf waters during 1991. Netherlands Journal of Sea Research 34: 45-58.

Wootton, R.J. 1979. Energy costs of egg production and environmental determinants of fecundity in teleosts fishes. In: P.J. Miller (Ed), Fish Phenology: Anabolic Adaptiveness of Teleost. pp: 133-159.


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