Ctenopharyngodon idella (Valenciennes, 1844)

Features
Simular Species
Biology
Maximum Size
Distribution
Interest to Fisheries
Current Status
Impacts
Recommendations
References

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Scientific Name: Ctenopharyngodon idella

Integrated Taxonomic Information System (ITIS): 163537

Other scientific names appearing in the literature of this species:

Common Name: Grass carp, white-amure

Distinguishing Features:

Soin (1963) and Soin and Sukhanova (1972) describe numerous differences between the eggs and larvae of C. idella and the black carp, Mylopharyngodon piceus Richardson, the silver carp, Hypopthalmichthys molitrix Valenciennes, and the bighead, Hypopthalmichhtys nobilis (Richardson).

Similar Species:

Taxonomy: Howes (1981) reviewed the phylogenetic position of this species within the Cyprinidae and assigned Ctenopharyngodon idella to the Squaliobarbine group of Asian cyprinds. He concluded it to be the sister-species of Mylopharyngodon piceus (Richardson, 1845) and that these two species together form the sister taxon of the two species of Squaliobarbus.

Biology:

What is known about the grass carp predominantly results from its usefulness in vegetation control (Shireman and Smith, 1983) and their importance as food (Bardach et al., 1972).

Soft aquatic vegetation preferred by this herbivorous species in its natural and introduced habitat includes filiform, Ziz's pondweed, pondweed, hornwort, spiked milfoil, duckweed and water thyme. However, in their absence C. idella will consume corser macrophytic species including reeds, reed sweet-grass, reed-mace, sedges, bulrushes, and horsetail Verigin et al., 1963; Stroganov, 1963; Nikol'skiya and Verigin, 1966; Bobrova, 1968; Yaroshenko, et al., 1970; Gurova, 1972). In high numbers the grass carp has been reported to have significant impact on macrophytes and can remove, essentially all aquatic vegetation. All vegetation can be eliminated, even in large aquatic systems such as the 8,100 ha Lake Conroe in Texas (Klussmann, et al. 1988; Maceina, et al., 1992).

Grass carp spawned in the Ili River in 1972 and 1973 from mid May through late June, when water temperatures reached 18.7° and 23.5° respectively, with peak spawning noted when temperatures reached 19.5-19.9°. Spawning commenced after a rise in water level and considerable increase in turbidity (Nezoliy and Mitrofanov, 1975). In the Amur River, spawning takes place in the summer (June-July), with eggs drifting downstream at temperatures of 17-19°C, primarily near the surface, to the beginning of August (Krykhtin and Gorbach, 1981). Water level also appears to be important, with peak spawning associated with rises in water level, usually commencing only during rises above 20 cm to greater than 2 m and ceasing when the level begins to lower (Kryhtin and Gorbach, 1981).

A 6 kg female from the Yangtse was reported to have more than 100,000 eggs (Chen and Lin, 1935), while others in the same drainage near Ichang had between 29,000 to 138,000 eggs (Nikol'skiy, 1956). Anishchenko (1939) noted 816,000 eggs in a 76cm 7.4 kg female. Chtang Yu-fan, et al. (1961) reported a 14.62 kg female bearing 960,000 eggs. Inaba et al. (1957) reported 485,000 eggs in a 88 cm, 7.135 kg female, and 800,000 eggs in a 81.5cm 12 kg fish, both from Japan. In India females between 2-6 kg were reported with fecundities of 82 eggs/g of body weight (Alikunhi, et al., 1963). In Malacca pond raised grass carp were found to bear between 2,000 and 37,000 eggs/kg of body weight (Hickling, 1967). In European latitudes fecundity ranged between an average 500,000-700,000 eggs in fish with an average weight of between 6-8 kg (Prikhod'ko and Nosal', 1963; Vinogradov, 1966; Aliyev, 1968; Vinogradov, et al., 1968; Vovk, 1968). In the Amur Basin found grass carp to bear between 237,00 to 1,687,000 eggs, with an average of 820,000 eggs in 7+ to 15+ year old fishes between 66-96 cm and 5.05-16.4 kg (Gorbach, 1972). Gorbach found fecundity is lowest in first time spawners and highest in large females that have spawned repeatedly.

This species is known to hybridize with Hypopthalmichthys molitrix (Verigin, Makeyeva, and Shubnikova, 1975; Marian and Krasznai, 1978).

Grass carp eggs may develop heterochronically, with an inverse relationship between development time of embryos in the egg capsule and time of hatching (Goryunova, 1971). Growth of juvenile carp in hatcheries was reported as inversely related to stocking density (Le Hoa, 1973). Condition factor for Amur River populations were reported as different between the sexes with males growing more quickly at small sizes and with females growing more quickly at large sizes (Gorbach, 1971).

Electophoretic analysis of muscle tissue has permitted differentiation of this species with the Silver Carp (Hypopthalmichthys molotrix and the Bighead Hypopthalmichthys nobilis, as well as among geographic variants and hybrids between the species (Truveller et al., 1973; Payusova and Tselikova, 1982).

Maximum Size:

This species grows to 125 cm in length (Page and Burr, 1991).

Distribution:

The grass carp is native to the middle and lower Amur River, the Sungari and Ussuri Rivers, and Lake Khanka, and to eastward flowing rivers of the People's Republic of China south to Guangzhou, Kwangtung Province (Courtenay et al., 1984).

Interest to Fisheries:

This species may play an indirect role in supporting fisheries of species that feed on phytoplankton by reducing macrophytic competitors of phytoplankton species (Charynev, 1980, 1984). Nikol'skiy and Aliyev (1974) and Aliyev (1976) reported that phytophagous species (Hypopthalmichthys molitrix and Hypopthalmichthys noblis accounted for 75-80% of the catch, or 45 kg/hectare of the Khauz Kahn Resevoir fishery, a figure Charynev (1984), argued was largely due to these species consuming phytoplankton that "bloomed" when macrophytes were consumed by grass carp. The current status of these fisheries is unknown. Maceina et al. (1992) reported changes in biotic diversity concomitant with reduction of areal coverage of submerged macrophytes from 44% to 0% in Lake Conroe. Densities of rotifers, cladocerans, and total zooplankton decreased approximately 1.5 years after macrophyte removal. Likewise, Lepomis spp. biomass declined, as reported by Bettoli, 1987). However, the abundance of Dorosoma petense, Menidia beryllina, Notropis venustus, and Pimephales vigilax increased (Betolli et al., 1990; Maceina et al., 1992).

Current Status of this Species in the Gulf of Mexico Ecosystem:

The spread of grass carp throughout the United States between 1963 and 1976 was summarized by Guillory and Gasaway, (1978), primarily through its use as a biological control of aquatic macrophytes, such as Hydrilla verticillata, Potamogeton illinoensis, Eleodea spp., Ceratophyllum dermersum, and Najas quadalupensis (Leslie et al., 1986; Shireman and Smith, 1983; Von Zon, 1979).

This species was recommended as a potential agent for control of aquatic "weeds" in (Swingle, 1957). Grass carp were first imported into the United States from Malaysia in 1963 (Pflieger, 1978) by the Fish Farming Experimental Station, Stuttgart, Arkansas and by Auburn University, Alabama (Stevenson, 1965). Fry from the grass carp held at the Fish Farming Experimental Station accidently escaped into the White River near Stuttgart, Arkansas in 1966 (Bailey, 1972). Grass carp were first reported outside of Arkansas in the Illinois portion of the Mississippi River in 1971 (Greenfield, 1973) and latter in the lower Mississippi (Conner et al., 1980).

Stanley, et al. (1978) noted that this species would likely reproduce widely throughout North America, once introduced. Despite efforts to limit its distribution in natural systems, its spread has been extremely rapid. In addition to the Mississippi, it is now established in Texas (Nobel et al., 1986), in the Ohio River (Jennings, 1989), and in the Missouri River system (Brown and Coon, 1991).

Because concern of unwanted introductions and undesirable impacts on natural vegetation, all 50 states have some restrictions on their use in weed control. Initial efforts to prevent unwanted introductions included creation of all female populations through gynogenesis (Stanley, 1976), through hormonal implants followed by mating sex-reversed males to normal females (Boney et al, 1984), or through gonadectomy. However, such monosexual populations remain fertile and chance introduction or production of males may lead to reproduction (Chourrout and Quillet, 1982). Godadectomy are not effective in stemming reproduction because the gonads are capable of rapid regeneration (Clippinger and Osborne, 1984; Underwood et al., 1986).

Discovery that F1 hybrids of intergeneric crosses between female C. idella and male Hypophthalmichtys nobilis were 100% triploid, led to the use of triploids in commercial weed control (Malone, 1984). However, subsequent commercial efforts to duplicate this finding produced only 67% triploidy (Allen and Wattendorf, 1987). Later, Cassani and Caton (1985) discovered that cold shocks could produce 50-100% triploidy with egg survival of less than 20%. Extending these findings, Cassani and Caton (1986) utilizing hydrostatic pressure treatments at 7000 to 8000 psi produced nearly 100% triploidy with 30% mortality. Bonar et al. (1984) noted that diploids could be differentiated from triploids on the basis of meristics and morphometrics, but only through use of multivariate methods. Consequently, enforcement of stocking regulations may be difficult without determination of ploidy through microscopic examination or the use of a Coulter Counter. The latter is a device that takes advantage of differential resistance of particles, in this case erythrocytes, of different size in passing through an orifice between two electrodes. Because triploids have larger erythrocytes and nuclei than diploids, it is possible to separate the two with nearly 100% accuracy (Beck and Biggers, 1983; Benfey et al, 1984; Johnson et al., 1984; Wattendorf, 1986). Triploid grass carp produce only rudimentary gonads (Doroshov, 1986). Allen et al. (1986) and Allen and Wattendorf (1987) noted that while it is possible for triploid carp to produce offspring through mating with diploid adults, the vast majority are sterile, probably because the sperm are aneuploid. Consequently, although reproductive they are functionally sterile, with only 0.00000012 of the gametes fertile.

Potential Impacts:

Grass carp were widely introduced widely in the European territories of the former USSR, Central Asia, and Kazakhstan both at the Amu Darya River (Aliyev, 1965) and the Balkhash-Ili basin. During 1963-1973 22237 adults and 6,309,500 young were introduced into natural lakes of the latter basin (Nezdoliy and Mitrofanov, 1975), the Kuban River (Motenkov, 1972), and the Lower Volga (Martino, 1974). They were introduced into Turkmenistan in 1958 and into the Kara Kum Canal in large numbers in 1960-1961 (Charyev, 1984). According to Charyev, this species prevents reeds, (Phragmites communis and reed-mace, Typha angustata from growing in water deeper than 50-60 cm in the Kara Kum, not only by reducing the number of new shoots growing from rootstock, but also actively consuming parts above the waterline. He further reported that it feeds extensively on water milfoil Myriophyllum spicatum, allowing older water crowfoot Ranunculus to become dominant, although young shoots of the latter species were consumed in entirety at other locations. In high numbers the grass carp has been reported to have significant impact on macrophytes and can remove, essentially all aquatic vegetation. Lake Conroe macrophytic species eradicated through consumption by grass carp, included hydrilla, Hydrilla verticillata, Eurasian watermilfoil, Myriophyllum spicatum, and coontail, Ceratophyllum demersum (Maceina et al., 1992). In lesser densities, their herbivory is more selective and has led to increased in "inedible" or toxic plants (Kogan, 1974; Vinogradov and Zolotova, 1974; Charyev, 1980; Charyev, 1984), thus reducing water filtration by macrophytes.

Allen and Wattendorf (1987) discussed use of the species in the US for aquatic plant control and concluded that its introduction is not a panacea for plant control, since they may total eliminate aquatic plants, thereby eradicating habitat for invertebrates and juvenile fishes to the detriment of sport fisheries or forage for waterfowl to the detriment of hunters. Webb et al. (1987, 1989) noted significant declines in gadwall, Anas stepera, American wigeon, Anas americana, and American coot Fulica americana after grass carp introduction.

Because abundant macrophytes affect water-column nutrient concentrations (Wetzel, 1975), alter trophic interactions (Body, 1971), and cause changes in algal abundance (Goulder, 1969; Canfield et al., 1984; and affect zooplankton communities (Richards et al., 1985), it is to be expected that introduction of grass carp are likely to reverse many such effects, thereby affecting community structure and phytoplankton and zooplankton abundance and size (Lizzaro, 1987; Maceina, et al., 1992). Taylor et al. (1984) report increased nutrient concentrations and phytoplankton abundance in ponds stocked with grass carp. However, Mitzner (1978) and Leslie et al, (1983) did not report an increase in nutrient or algal levels in lakes whose vegetation was controlled through use of grass carp.

Allen and Wattendorf (1987) suggested that their use may be better adapted to situations in which total plant elimination is desired, such as golf course ponds, small urban lakes, irrigation canals, or areas where plants would otherwise be eliminated through the use of herbicides. Bain (1990; 1993) notes, however, that because they make extensive spawning migrations and congregate in turbulent habitats associated with rapids, falls, and dams, grass carp are capable of moving well beyond areas intended for plant control within a single season. Migrations of up to 1700 km are known for this species (Guilory and Gasaway, 1978).

This species has been reported as introduced to control the malarial mosquito Anopheles pulcherrimus and Culex species in the Kara Kum Canal of the former Soviet Union (Aliyev and Bessmertnaya, 1968). It was hypothesized by these authors that mosquito larvae associated with vegetation were consumed through herbivory by this species. These authors further noted that local Gambusia disappeared after such introduction, as well as an unspecified number of "small worthless fishes", which were presumably deprived of refuge and thus made more visible to predators.

Recommendations:

References:

Alikunhi, K.H., K.K, Sukumaranan, Parameswaran. 1962. Induced spawning of the Chinese grass carp, Ctenopharyngodon Idellus (C. et V.) and the Sllver carp, Hypophthalmicthys molitrix (C. et V.), in ponds at Cultack, India. Proc. Indo-Pacif.Fich. Conc., Vol. 10, No. 2.

Aliyev, D. S. 1965. The reproduction of the grass carp Ctenopharyngodon idella and of the silver and bighead carps Hypophthalmichthys molitrix and Aristichthys nobilis established in the Amur Darya basin. Journal of Ichthyology. 5(4):37.

Aliyev, D. S. 1968. Fundamental questions of the biological technique for the artificial propagation of phytophagous fish under productive conditions. In: Novyye issledovaniya po ekologii i razvedeniyu rastitel'noyadnykh ryb. Moscow, Nauka Press.

Aliyev, D. S. 1976. The role of phytophagous fishes in the restoration of ichthyofauna and biological improvement of bodies of water. Journal of Ichthyology, 16(2):247-262.

Aliyev, D. S., and R. Ye. Bessmertnaya. 1968. Use of the grass carp, [Ctenopharyngodon idella (Val.)] to control the larvae of blood-sucking mosquitoes. Problems in Ichthyology, 8(2):319-321.

Allen, J., S.K., R.G. Thiery, N.T. Hagstrom. 1986. Cytological evaluation of the likelihood that triploid grass carp will reproduce. Transactions of the American Fisheries Society 115(6):841-848.

Anishchenko 1939. On The Acclimation Of Amur Fish In The European Areas Of The USSR. Rybn. Khoz. No. 5.

Bailey, W. M. 1972. Arkansas's evaluation of the desirability of introducing the white amur (Ctenopharyngodon idella) for control of aquatic weeds. Arkansas Game and Fish Commission, mimeo.rep. 59 pp.

Bain, M. B. 1993. Assessing impacts of introduced aquatic species: grass carp in large systems. Environmental management, vol. 17(2):211-224.

Bardach, J. E., J. H. Ryther, and W. O. McLarney. 1972. Aquaculture: the farming and husbandry of freshwater and marine organisms. Wiley-Interscience, New York, 868 pp.

Barnett, B. S., and R. W. Schneider. 1974. Fish populations in dense, submersed plant communities. Hyacinth Control Journal 12:12-14.

Beaty, P.R., R.G. Thiery, N.T. Hagstrom, M.R. Mizumoto. 1984. Impact of hybrid amur in two California irrigation systems: 1983 Progress Report. Coachella Valley Water District, Coachella, Ca. 59 pp.

Beck, M.L. and C.J. Biggers. 1983. Erythrocyte measurements in diploid and triploid Ctenopharyngodon idella x Hypophthalmichthys nobilis hybrids. Journal of Fish Biology 22:497-502.

Benfey, T. J., A. M. Sutterlin, and R. J. Thompson. 1984. Use of erythrocyte mesurements to identify triploid salmonids. Can. J. Fish. Aquatic Sciences. 41:980-984.

Berry, P. Y. and M. P. Lowe. 1970. Comparative studies on some aspects of the morphology and histology of Ctenopharyngodon idellus, Aristichthys nobilis and their hybrid (Cyprinidae). Copeia 1970(4):708-726.

Bettoli, P. W. 1987. The restructuring of a forage fish community following large-scale aquatic vegetation control. Ph.D. Dissertation. Texas A&M University. College Station, TX.

Bettoli, P. W., J. Morris, and R. L. Noble. 1990. Changes in the abundance of two silversides species afer aquatic vegetation removal. Trans. Am. Fish. soc. 120:90-97.

Bobrova, Y. P. 1968. The Feeding And Growth Of The Grass Carp Under Pond Fish Farming Condidtions In The Central Zone Of The RSFSR. In: Novyye Isseldovaniya Po Ekologii I Razvendeniyu Rastitel'noyadnykh Ryb. Moscow, Nauka Press.

Bonar, S. A., G. L. Thomas, G. B. Pauley, A. Unthank. 1984. An evaluation of ploidy separation techniques for grass carp (Ctenopharyngodon idella), a potential biological control of aquatic macrophytes in Washington state. Pages 158-164 in Proc. of the 19th Annual Mtg. Aquatic Plants Control Res. Program, Army Corps of Engineers, 26-29 Nov 1984, Gavelston, Tx.

Boney, S. E., W. L. Shelton, S. L. Yang, and L. O. Wilken. 1984. Sex reversal and breeding of grass carp. Transactions of the American Fisheries Society 113:348-353.

Buck, H. 1979. Optimism swells with the possibility of a sterile hybrid grass carp. Fisheries 4(5):31.

Cassani, J. R. and W. E. Caton. 1986. Efficient production of triploid grass carp (Ctenopharyngodon idella utilizing hydrostatic pressure. Aquaculture, 55:43-50.

Charyev, R. 1980. The grass carp and the phenomenon of succession in bodies of water. Ecology, No. 4, 93-94.

Charyev, R. 1984. Some consequences of the introduction and acclimatization of grass carp, Ctenopharyngodon idella (Cyprinidae), in the Kara Kum Canal. Journal of Ichthyology 24(3):1-8.

Chen, C. S. and S. V. Lin. 1935. The fish fry industry of china. Bull Chekiang Province Fisheries Experiment Station. Tinghai, 1(4).

Chourrout, D. and E. Quillet. 1982. Induced gynogenesis in the rainbow trout: sex and survial of progenies. Production of all triploid populations. Theor. Appl. Genet. 63:201-205.

Chtang Yu-fan, Cheng Chieh, Wang Chiang-Chi, and Liu Tsao-fai. 1961. Khozyaystvo Vnutrennikh Vod Kitaya (The Fish Industry Of Chinese Inland Waters). Peking.

Clippinger, D. and J. A. Osborne. 1984. Surgical sterilization of grass carp, a nice idea. Aquatics 6:9-10.

Conner, J. V., R. P. Gallagher, and M. F. Chatry. 1980. Larval evidence for natural reproduction of the grass carp (Ctenopharyngodon idella) in the lower Mississippi River. Pages 1-19 in , L. A. Fuiman, editor. Proceedings of the fourth annual larval fish conference held 27-28 February 1980 in Oxford, Miss. U.S. Fish and Wildlife Service, Ann Arbor, Mich. FWS/OBS 80/43.

Courtenay, W. R., Jr., D. A. Hensley, J. N. Taylor, and J. A. McCann. 1984. Distribution of exotic fishes in the continental United States. Pages 41-77 in W. R. Courtenay, Jr., and J. R. Stauffer, Jr., editors. Distribution, biology and management of exotic fishes. Johns Hopkins University Press, Baltimore, MD.

Doroshov, J. 1986. Comparative gametogenesis in diploid and triploid grass carp. Meeting of World Mariculture Soc., Reno, NV. January 1986 (abstract).

Gorbach, E.I. 1971. Condition and fatness of the grass carp (Ctenopharyngodon idella (Valenciennes)) in the Amur Basin. Journal of Ichthyology 11:880-889.

Gorbach, E.I. 1972. Fecundity of the grass carp (Ctenopharyngodon idella (Val.)) in the Amur Basin. Journal of Ichthyology 12:616-625.

Goryunova, A.I. 1971. Variability In The Rate Of Embryogensis Of The Grass Carp (Ctenopharyngodon Idella (Val.)). Journal Of Ichthyology 11(1):44-48.

Greenfield, D. W. 1973. An Evaluation Of The Advisability Of The Release Of Grass Carp, Ctenopharyngodon Idella, Into Natural Waters Of The United States. Trans. Ill. State Acad. Sci. 66(1-2):47-53.

Guillory, V., and R. D. Gasaway. 1978. Zoogeography of the grass carp in the United States. Transactions of the American Fisheries Society 107(1):105-112.

Gurova, L. A. 1972. The feeding and growth of phytophagous fishes in the ponds of Chita power station. Zap. Zabaykal'sk. fil. Geogr. o-va SSR, No. 62.

Hickling, C. F. 1967. On the biology of a herbivorous fish the white amur or grass carp, Ctenopharyngodon idella Val. Proc. R. Soc. Edinburgh Sect. B 70(1):62-81.

Howes, G. 1981. Anatomy and phylogeny of the Chinese major carps Ctenopharyngodon Steind., 1866 and Hypophthalmichthys Blkr., 1860. Bulletin of the British Museum (Natural History) Zoology Series 41(1):1-52.

Inaba, D., M. Nomura, and M. Nakamura. 1957. Preliminary report on the spawning of the grass carp and silver carp in the Tone River, Japan and the development of their eggs. Journal of Tokyo University of Fisheries 43:81-101.

Johnson, O. W., P. S. Rabinovitch, and F. M. Utter. 1984. Comparison of the reliability of a coulter Counter with a flow cytometer in determining ploidy leveles of Pacific salmon. Aquaculture 43:99-103.

Klussmann, W., R. Noble, R. Martyn, W. Clark, R. Betsill, P. Bettoli, M. Cichra, J. Campbell. 1988. Control of aquatic macrophytes by grass carp in Lake Conroe, Texas, and the effect on the reservoir system. Texas Agriculture Experiment Station MP-1664, College Station, Texas.

Kogan, S. I. 1974. Overgrowing of the Kara Kum Canal and some after effects of introducing the white amur into water bodies. Gidrobiol. Zh. 10(2):110-115 (in Russian).

Lazzaro, L. 1987. A review of planktivorous fishes: Their evolution, feeding, behavior, selectivities, and impacts. Hydrobiologia. 146:97-167.

Le Hoa, D.T. 1973. Variability of juvenile grass carp (Ctenopharyngodon idella (Val.)) and pond carp (Cyprinus carpio (L.)) reared at a hatchery in the southern Ukraine. Journal of ichthyology 13(2):305-309.

Leslie, A. J., Jr., J.M. Van Dyke, R.S. Hestand III, B.Z. Thompson. 1987. Management Of Aquatic Plants In Multiuse Lakes With Grass Carp (Ctenopharyngodon Idella). In: Redfield, Ed. Lake And Reservoir Management, Vol. 3, Proc. Of N.A. Lake Management Soc., Portland, Oregon.

Leslie, A. J., Jr., L.E. Nall, J.M. VanDyke. 1983. Effects of Vegetation Control by grass carp on selected water quality variables in four Florida lakes. Transactions of the American fisheries Society 112:777-787.

Maeceina, M. J., M. F. Cichra, R. K. Betsill, and P. W. Bettoli. 1992. Limnological Changes in a Large Reservoir Following Vegetation Removal by Grass Carp. J. Fresh. Ecology 7(1):81-95.

Malone, J.M. 1984. Triploid white amur. Fisheries (Bethesda)9(2):36.

Marian, T. and Z. Krasznai. 1978. Karyological investigations on Ctenopharyngodon idella and Hypophthalmichthys nobilis and their cross breeding. Aquacultura Hungarica (Szarvas)1:44-50.

Martino, K. V. 1974. Natural reproduction of the grass carp in the lower Volga area. Hydrobiological journal 10(1):76-78.

Mitzner, L. 1980. Evaluation Of Biological Control Of Nuisance Aquatic Vegetation By Grass Carp. Transactions Of The American Fisheries Society 107:135-145.

Motenkov, Y. M. 1972. The results of the establishment of grass carp, silver carp and bighead carp in the waters of the Kuban River basin and the prospects for the commercial utilization of these fish. Thesis, Leningrad.

Nezdolii, V.K., V.P. Mitrofanov. 1975. Natural reproduction of the grass carp, Ctenopharyngodon idella, in the Ili River. Journal of Ichthyology 15:927-933.

Nikol'skiy, G. V. 1956. Fishes of the Amur Basin. Moscow Academy of Science U.S.S.R. 551 pp.

Nikol'skiy, G. V. and B. V. Verigin. 1966. The main biological features of the grass carp and the silver carp and their acclimatization in the waters of the country. In: Rastitel'noyadnyye ryby. Moscow, Pischevaya promyshelnnost.

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Noble, R.L., P.W. Bettoli, R. Betsill. 1986. Considerations for the use of grass carp in large open systems. Lake and Reservoir Management 2:46-48.

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Payusova, A. N. And T. N. Tselikova. 1982. Differentiation Of Populations Of Grass Carp, Ctenopharyngodon Idella, Silver Carp, Hypophthalmichthys Molitrix, and Bighead, Aristichthys Nobilis, From Electrophoresis Myogens. Journal Of Ichthyology. (20):23-30.

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Underwood, J. L., R. S. Hestand, and B. Z. Thompson. 1986. Gonad regeneration in grass carp following bilateral gonadectomy. Prog. Fish-Cult. 48-54-56.

Verigin, B. V., A. P. Makeyeva, and N. G. Shubnikova. 1975. The morphology of the underyearling hybrids of the bighead Aristichthys nobilis and the grass carp Ctenopharyngodon idella. Journal of Ichthyology 15(2):226-231.

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