Clupeonella cultriventris (Black Sea kilka)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Water Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Vectors
- Impact Summary
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Clupeonella cultriventris (Nordmann, 1840)
Preferred Common Name
- Black Sea kilka
Other Scientific Names
- Clupea cultriventris Nordmann, 1840
- Clupea cultriventris tscharchalensis Borodin, 1896
- Clupea delicatula Nordmann, 1840
- Clupeonella caspia Svetovidov, 1941
- Clupeonella cultiventris caspia Svetovidov, 1941
- Clupeonella delicatula Nordmann, 1840
- Clupeonella delicatula caspia Svetovidov, 1941
- Clupeonella delicatula cultriventris Nordmann, 1840
- Clupeonella tscharchalensis Borodin, 1896
International Common Names
- English: Azov Sea kilka; Caspian Sea kilka; kilka; Ponto-Caspian kilka
- Spanish: espadín del Mar Caspio; espadín del Mar de Azov; espadín del Mar Negro
- French: clupeonelle
- Russian: chernomorsko-kaspiyskaya kilka (tulka); obyknovennaya tyulka, obyknovennaya kilka
- Portuguese: espadilha do Cáspio; espadilha-do-Mar-Negro
Local Common Names
- Bulgaria: tzatza
- Czech Republic: kilka kaspická
- Denmark: kaspisk sild
- Germany: Kaspische Sprotte
- Iran: kilka-e-maamooli
- Italy: papaline del Caspio
- Kazakhstan: kilkas tyulka lar
- Netherlands: zwarte-zeesprot
- Poland: kilka kaspijska
- Romania: gingirica
- Sweden: kaspisk skarpsill
- Turkey: kilka baligi
- Turkmenistan: adaty kulke balyk
- Ukraine: tyulka
Summary of InvasivenessTop of page
C. cultriventris, a Ponto-Caspian endemic, inhabits the northwestern parts of the Black Sea, the Sea of Azov and the Caspian Sea, also most rivers of the area, and some lakes in Bulgaria, Romania and Turkey (Stojanov, 1963; Whitehead, 1985; Aseinova, 1989, 1992; Alexandrov et al., 2008).
C. cultriventris is considered a medium to high risk invader. It was identified as having a high probability of invasion if introduced to the Great Lakes via ballast water (Ricciardi and Rasmussen, 1998; Kolar and Lodge, 2002,) and the Baltic Sea via natural and artificial waterways (Panov et al., 2007). However, Slynko (Institute for Biology of Inland Waters, Russia, personal communication, 2011) notes that the probability of invasion via ballast waters is low as the eggs and young are easily damaged. Dispersion of C. cultriventris is still limited within the Black and Caspian Sea basin although C. cultriventris has already been recorded in the River Pripyat (Belarus), very close to Baltic Sea basin (Semenchenko et al., 2009). Since the 1980s an abundance of Mnemiopsisleidyi (a severe competitor to plankton-feeding fish) in the Black and Azov seas and later the Caspian Sea led to a sharp decline in stocks in the Black and Asov seas (Aseinova, 1989, 1992; Zaitsev et al., 1997), therefore lowering risk of introduction via ballast water. C. cultriventris stocks did not decline in the Caspian Sea due to their high plasticity and capacity to migrate for feeding to the areas with low salinity, which are not available for M. leidyi (Shiganova, 2011). Contrary to other countries, in Bulgaria and Romania C. cultriventris is considered as a vulnerable species (Schiemer et al., 2004).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Chordata
- Subphylum: Vertebrata
- Class: Actinopterygii
- Order: Clupeiformes
- Family: Clupeidae
- Genus: Clupeonella
- Species: Clupeonella cultriventris
Notes on Taxonomy and NomenclatureTop of page
DescriptionTop of page
Max. length: 14.5 cm TL; max. reported age: 5 years.
Body comparatively deep, maximum depth 15.5-23.5% of TL, 19.7% average. Head moderate, average 21% of TL. Mean eye diameter 25% of head length. Lower jaw slightly extended. Ventral scutes well developed. Pectorals and pelvics long, pectorals 17.7-21.2% (average 12.7%) of TL, pelvics 11.7-13.8% (average 12.7%) of TL. Gill-rakers: 51-62 (average 60). Vertebrae 40-43 (average 41). The back and the upper part of head vary in colour from light green to blue-green, abdomen silvery white or golden yellow (Whitehead, 1985; Aseinova, 1989, 1992; Hoestlandt, 1991).
DistributionTop of page
Origin: A Caspian autochthon.
World distribution: A Ponto-Caspian endemic species.
Climate temperate: 60°N - 36°N, 27°E - 56°E.
Eurasia: Black Sea (mainly northwestern parts), Sea of Azov and Caspian Sea, also most rivers of the area, it has entered the Volga, Ural, Dneiper, Dneister and Terek rivers (Whitehead, 1985; Aseinova, 1989, 1992). In Bulgaria: the Lakes Shabla, Varna, Burgas, Palaeostomi; the Bay of Varna; in Romania: the bay of Feodosiya, Tasaul Lake; in Turkey, Lake Apolyont (Stojanov, 1963; Banarescu, 1964; Alexandrov et al., 2008). The trend of dispersion is northwards in Volga and Dnieper rivers – it was recently found in Pripyat River (Dnieper tributary) and Ivanovo reservoir in Volga River (Leppäkoski et al., 2002; Semenchenko et al., 2009).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Mediterranean and Black Sea||Widespread||Native||Not invasive||Whitehead, 1985; Zaitsev and Alexandrov, 1997||Only Black Sea|
|Azerbaijan||Widespread||Native||Not invasive||Whitehead, 1985|
|Georgia (Republic of)||Present||Native||Not invasive||Svetovidov, 1964|
|Iran||Widespread||Native||Not invasive||Whitehead, 1985|
|Kazakhstan||Widespread||Native||Not invasive||Whitehead, 1985|
|Turkey||Present||Native||Not invasive||Bilecenoglu et al., 2002|
|Turkmenistan||Widespread||Native||Not invasive||Whitehead, 1984|
|Belarus||Present, few occurrences||Introduced||1986||Not invasive||Semenchenko et al., 2009||River Pripyat|
|Bulgaria||Present||Native||Not invasive||Whitehead, 1985|
|Romania||Present||Native||Not invasive||Whitehead, 1985|
|Russian Federation||Present||Present based on regional distribution.|
|-Central Russia||Localised||Introduced||Invasive||Leppäkoski et al., 2002; Stepanov and Kiyashko, 2008; Slynko et al., 2011||Kuybyshev, Rybinsk, Gorki, Ivanovo reservoirs (Volga Basin)|
|-Southern Russia||Widespread||Native||Not invasive||Reshetnikov et al., 1997|
|Ukraine||Widespread||Native||Not invasive||Whitehead, 1985; Ryabov, 2002||Invasive in all parts of the Dnieper above the former thresholds, i.e. in reservoirs from Zaporozhskoye to Kievskoye inclusive (Y Slynko, Institute for Biology of Inland Waters, Russia, pers. comm., 2011)|
History of Introduction and SpreadTop of page
In the Volga basin: C. cultriventris was established by the mid-1960s in the Kuybyshev Reservoir, by mid-1970s it was found in the Gorki Reservoir, in the 1990s it was established in the Rybinsk Reservoir, in 2000 it was found in Ivanovo Reservoir and it is now found up to the Beloye Lake (Kiyashko et al., 2006).
In Dnieper River basin C. cultriventris was recently found to be present in all reservoirs up to the River Pripyat (Semenchenko et al., 2009).
Risk of IntroductionTop of page
C. cultriventris is considered a medium to high risk invader. C. cultriventris was identified as having a high probability of invasion if introduced to the Great Lakes via ballast water (Ricciardi and Rasmussen 1998; Kolar and Lodge, 2002). Since the 1980s an outbreak of Mnemiopsis leidyi (a severe competitor to plankton-feeding fish) in the Black and Azov Sea and later the Caspian Sea led to a sharp decline in stocks (Aseinova, 1989, 1992; Zaitsev et al., 1997), therefore lowering risk of introduction via ballast water. Although most of the animals that have invaded the Great Lakes since 1985 are Ponto-Caspian in origin (Ricciardi and MacIsaac, 2000), C. cultriventris absence in the Baltic sea region, usually the first region to invade for almost all well known Ponto-Caspian invaders (Ojaveer et al., 2002), indicates that likelihood of introduction remains relatively low. However C. cultriventris dispersal to the upper reaches of Dnieper and Volga rivers (both connected with the Baltic Sea basin via channels) indicates that dispersal to Baltic Sea is very likely (Kiyashko et al., 2006; Semenchenko et al., 2009).
HabitatTop of page
C. cultriventris is a euryhaline species. It is found in coastal waters, lagoons and lakes, estuaries and lower reaches of large rivers with a salinity up to 13 ‰. The biggest concentrations are recorded in zones with salinity ranging from 3 to 7‰. Increased salinity (up to 12‰) during low water periods is supposed to be one of the main factors which affect kilka survival. The most intensive spawning occurs at salinity values 2-4‰. It is absent from pure seawater.
It is a eurythermic species, occurring at surface temperature ranges of 2.6 to 26°C. The optimal temperature for this species is 16-22°C, and the optimal spawning temperature is 14-20°C (Y Slynko, Institute for Biology of Inland Waters, Russia, personal communication, 2011).
C. cultriventris is a eurybathic and oxyphilic species.
It is migratory between winter or autumn feeding and summer spawning grounds, populations in the Black Sea are anadromous and it spawns in rivers. Populations in the Caspian and Azov seas are partially anadromous (it spawns in the sea as well in rivers) (Whitehead, 1985; Aseinova, 1989, 1992).
Habitat ListTop of page
|Inland saline areas||Secondary/tolerated habitat||Natural|
|Coastal areas||Principal habitat||Natural|
|Irrigation channels||Secondary/tolerated habitat||Natural|
|Rivers / streams||Principal habitat||Natural|
Biology and EcologyTop of page
The total mitochondrial genome of C. cultriventris has been sequenced (http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=501719).
C. cultriventris breeds in the late spring to early summer mainly in the brackish Taganrog Bay of the Sea of Azov (peak in May), and from about mid May in the lower reaches of rivers (Dneiper, Dneister). In the Caspian Sea, C. cultriventris starts spawning in April and continues until late June, its peak is observed in the first 10 days of May at a water temperature of 14-19°C, and depth of 1-6 m. In May, spawning takes place in the coastal areas, whilst in June it mostly takes place in waters 5-6 m deep. In the southern part of the Caspian Sea spawning starts in February (Aseinova, 1989, 1992).
Fecundity varies from 9.5 to 60 thousand eggs (31.2 thousand eggs on average). For more information see Osipov and Kiyashko (2006).
Life history stages
C. cultriventris is a batch spawning species. It produces small-sized pelagic eggs 1 mm in diameter, with a large oil drop occupying about 1/3 of the egg diameter, with a considerable perivitelline space. Eggs are retained in the water column owing to the oil drop. Depending on the water temperature hatching occurs 27-30 hours after spawning. Larvae are 1.3-1.8 mm TL with a large yolk sac. By September, fingerlings reach 50-55 mm TL (Aseinova, 1989, 1992).
Early ontogenesis proceeds in surface layers above the depths 1-7 m, where the most abrupt fluctuations of water temperature and surge occur. Stormy conditions at sea decrease the survival of embryos and larvae due to hydrodynamic wave impacts. The best survival of embryos and larvae is restricted to temperature ranges of 14-19°C. Eggs and larvae are found at comparatively wide salinity range (0.02 - 15‰) (Aseinova, 1989, 1992).
C. cultriventris matures at an earlier age than other related species. Most individuals mature at the age of 1 year, 45-70 mm TL. The population structure is characterized by a high abundance of recruitment and low abundance of the remainder (Aseinova, 1989, 1992).
The most rapid growth rate is recorded in the first month of life. During this period, sizes increase from 1.3-1.8 mm (newly hatched larvae) to 30.4 mm, i.e. almost 20-fold. During the second month, the total (absolute) increment comprises 12.1 mm, and 9.8 mm over the third month. In July, average size is 30.4 mm in the west, and 28.6 mm in the east; in August this is 42.5 mm and 40.9 mm, and in September this is 52.3 mm and 51.4 mm, respectively. Males grow slower than females; the most significant growth discrepancy is observed at the age of 2-3 years (Aseinova, 1989, 1992).
For more information see Kiyashko et al. (2006) and Osipov and Kiyashko (2008).
Heterotrophic (live organic substances).
Preying and selected capture of food items.
C. cultriventris is a euryphagous species. Their food spectrum contains copepods, cladocerans, rotifers, planktonic stages of Balanus and molluscs, crayfish and other components.
Quantitative characteristics of feeding
The daily diet of comprises 4.95–5.35% of body weight. Average daily food consumption (per specimen) is 0.163 g (Aseinova, 1989, 1992).
ClimateTop of page
|BS - Steppe climate||Tolerated||> 430mm and < 860mm annual precipitation|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Df - Continental climate, wet all year||Tolerated||Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Salinity (part per thousand)||3-7||Optimum||1-13 tolerated (Aseinova, 1989, 1992)|
|Water temperature (ºC temperature)||18||20||Optimum||2-26 tolerated (Aseinova, 1989, 1992)|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Acipenser persicus||Predator||All Stages||not specific||Moghadam et al., 2009|
|Acipenser stellatus||Predator||All Stages||not specific||Moghadam et al., 2009|
|Esox lucius||Predator||All Stages||not specific||Stepanov and Kiyashko, 2008|
|Lota lota||Predator||All Stages||not specific||Stepanov and Kiyashko, 2008|
|Perca fluviatilis||Predator||All Stages||not specific||Stepanov and Kiyashko, 2008|
|Sander lucioperca||Predator||All Stages||not specific||Stepanov and Kiyashko, 2008|
|Sander volgensis||Predator||All Stages||not specific||Stepanov and Kiyashko, 2008|
Notes on Natural EnemiesTop of page
Natural enemies of C. cultriventris include larger marine, brackish and freshwater fish, all piscivorous birds and mammals (seals, otters, bottlenose and common dolphins and common porpoise).
Means of Movement and DispersalTop of page
Natural Dispersal (Non-Biotic)
Possible as a contaminant of ballast water, but no specific cases are cited.
Pathway VectorsTop of page
Impact SummaryTop of page
|Economic/livelihood||Positive and negative|
|Environment (generally)||Positive and negative|
ImpactTop of page
C. cultriventris occupies a significant position in the brackish and freshwater ecosystems; it is both a basic consumer of zooplankton, and also a producer for predatory marine fish and seals.
It consumes the same zooplankton as young specimens of aboriginal fish species. In some cases feeding similarity exceeds 50% (Kiyashko et al., 2007), therefore high densities of C. cultriventris could alter growth of young aboriginal fish species.
About 60% of C. cultriventris stock is consumed by predators (Aseinova, 2003). In newly established areas C. cultriventris becomes a very important food item for almost all predatory fishes (Stepanov and Kiyashko, 2008).
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Tolerant of shade
- Highly mobile locally
- Fast growing
- Has high reproductive potential
- Altered trophic level
- Damaged ecosystem services
- Competition - monopolizing resources
- Rapid growth
- Difficult to identify/detect as a commodity contaminant
- Difficult/costly to control
UsesTop of page
C. cultriventris is an important commercial fish in the Azov, Black and Caspian seas (Aseinova, 2003). They are eaten fresh, salted, smoked, or tinned. In the former USSR they were an important part of the diet (Svetovidov, 1964).
Salted and tinned C. cultriventris in southern Ukraine and southern Russia are traditional foods, especially as part of Lent diets.
C. cultriventris could severely modify nutrient cycling in reservoirs through its feeding on zooplankton, which in cases of drinking-water reservoirs could have adverse implications for environmental services.
Uses ListTop of page
- Research model
- Sociocultural value
Detection and InspectionTop of page
C. cultriventris are small silvery fishes, elongate and somewhat compressed; belly with a distinct keel of scutes. Lower jaw projecting; gillrakers 41-62 on lower part of first arch. Pelvic fin origin below about middle of dorsal fin base; last two anal fin rays enlarged; no wing-like alar scales at base of caudal fin.
Similarities to Other Species/ConditionsTop of page
Clupeonella engrauliformis can be distinguished from C. cultriventris as the back and top of its head is dark violet whereas in C. cultriventris the back and the upper part of head varies in colour from light green to blue-green.
Prevention and ControlTop of page
Early warning systems
The list server L-ALIENS provides an international forum for announcing recent discoveries of non-native species. A similar, but peer-reviewed, function for non-native aquatic organisms is provided by the international journal, Aquatic Invasions (http://www.aquaticinvasions.net/), which was set up as part of the EC Integrated Project No. 506675 ‘Assessing LArge scale environmental Risks for biodiversity with tested Methods (ALARM)’.
This is established at the national level.
Other zooplankton eaters could effectively control populations of C. cultriventris. Since the 1980s an outbreak of Mnemiopsisleidyi (a severe competitor to plankton-feeding fish) in the Black and Azov seas and later in the Caspian Sea led to a sharp decline in stocks of C. cultriventris (Aseinova, 1989, 1992; Zaitsev and Mamaev, 1997). Predator fishes also provide good biological control as C. cultriventris are a very important food item for almost all predatory fishes in newly naturalized water bodies (Stepanov and Kiyashko, 2008).
The main risk of further dispersion is via ballast water, therefore management and control of ballast water is very important. C. cultriventris tolerates salinity up to 13‰, so ballast water change at sea could control further dispersion.
There are no known forms of biological control specific to C. cultriventris.
The only effective method of fish eradication is the application of rotenone, a pesticide that is also toxic to non-target native species.
Monitoring and Surveillance
Hydroacoustic methods and sampling.
Gaps in Knowledge/Research NeedsTop of page
Gaps in knowledge include information on containment/zoning and control of this species. There are only a few sources of information on the impacts of C. cultriventris outside its native range (Rybinsk Reservoir (Volga basin) only).
ReferencesTop of page
Alexandrov ML; Cernisencu I; Bloesch J, 2008. History and concepts of sustainable fishery in Tasaul Lake, Romania. GeoEcoMarina [Proceedings of the Swiss-Romanian Research Programme on Environmental Science & Technology.], 14:61-71. http://www.geoecomar.ro/publications/geoecomarina14.pdf
Khal'ko VV; Kiyashko VI; Khal'lko NA, 2009. Circadian Rhythmic Changes in the Lipid Composition of Muscles in Kilka Clupeonella cultriventris (Clupeidae, Clupeiformes) during Natural Periodicity of Feeding in the Feeding and Spawning Periods in the Rybinsk Reservoir. Journal of Ichthyology, 49(7):527-539.
Kiyashko VI; Osipov VV; Slyns'ko YV, 2006. Size-age characteristics and population structure of Clupeonella cultriventris at its naturalization in the Rybinsk Reservoir. Voprosy Ikhtiologii, 46(1):68-76.
Moghadam H; Tavakoli K; Pazhand M; Roufchahi ZA; Choubian R; Parand Avar FH, 2009. Summer and winter feeding behaviour in Acipenser persicus and Acipenser stellatus in the south Caspian Sea. Iranian Scientific Fisheries Journal, 18(2):13-26.
Nordmann A, 1840. [English title not available]. (Observations sur la fauna pontique.) In: Voyage dans la Russie méridionale et la Crimée. Voyage dans la Russie méridionale et la Crimée, II [ed. by Démidoff, A. de]. 353-635.
Ojaveer H; Leppakoski E; Olenin S; Ricciardi A, 2002. Ecological impacts of alien species in the Baltic Sea and in the Great Lakes: an inter-ecosystem comparison. In: Invasive Aquatic Species of Europe: Distributions, Impacts, and Management [ed. by Leppäkoski, E. \Olenin, S. \Gollasch, S.]. Kluwer Scientific Publishers.
Osipov VV; Kiyashko VI, 2006. Specific Features of the Reproduction of Clupeonella cultrivensis (Clupeiformes, Clupeoidei) at the Introduction into Freshwater Water Bodies. Journal of Ichthyology, 46(7):552-554.
Osipov VV; Kiyashko VI, 2008. Method of Age Determination of Ponto-Caspian Kilka Clupeonella cultriventris (Clupeiformes, Clupeoidei) Using Scales and Otoliths. Journal of Ichthyology, 48(8):637-643.
Panov VE; Dgebuadze YY; Shiganova TA; Filippov AF; Minchin D, 2007. A risk assessment of biological invasions in the inland waterways of Europe: the Northern Invasion Corridor case study. In: Biological invaders in inland waters: Profiles, distribution, and threats [ed. by Gherardi, F.]. Heidelberg, Germany: Springer, 639-656.
Reshetnikov YS; Bogutskaya N; Vasil'eva E; Dorofeeva E; Naseka A; Popova O; Savvaitova K; Sideleva V; Sokolov L, 1997. An annotated check-list of the freshwater fishes of Russia. Journal of Ichthyology, 37:687-736.
Ricciardi A; Rasmussen JB, 1998. Predicting the identity and impact of future biological invaders: a priority for aquatic resource management. Canadian Journal of Fisheries and Aquatic Sciences, 55(7):1759-1765.
Ryabov IN, 2002. Long-term observation of radioactivity contamination in fish around Chernobyl. In: Accident and Social Activities to Assist the Sufferers by the Accident [ed. by Imanaka, T.]. Research Reactor Institute, Kyoto University, 114-124.
Schiemer F; Holcik J; Keckeis H; Staras M, 2004. Ecological status and problems of the Danube river and its fish fauna: a review. In: Proceedings of the Second International Symposium on the Management of Large Rivers for Fisheries Volume I [ed. by Welcomme, R. L. \Petr, T.]. Bangkok, Thailand: FAO Regional Office for Asia and the Pacific, 273-299.
Semenchenko VP; Rizevsky VK; Mastitsky SE; Vezhnovets VV; Pluta MV; Razlutsky VI; Laenko T, 2009. Checklist of aquatic alien species established in large river basins of Belarus. Aquatic Invasions, 4(2):337-347. http://www.aquaticinvasions.ru/2009/AI_2009_4_2_Semenchenko_etal.pdf
Shiganova T, 2011. Non-native species in the Southern seas of Eurasia. Scientific World, 580 pp.
Slynko UV; Karabanov DP; Stolbunova VV, 2010. Genetic Analysis of the Intraspecific Structure of Kilka Clupeonella cultiventris (Nordmann, 1840) (Actinopterigii: Clupeidae). Doklady Biological Sciences, 433:261-263.
Slynko YuV; Dgebuadze YuYu; Novitskiy RA; Kchristov OA, 2011. Invasions of alien fishes in the basins of the largest rivers of the Ponto-Caspian Basin: composition, vectors, invasion routes, and rates. Russian Journal of Biological Invasions, 2(1):49-59. http://www.springerlink.com/content/f4514123l3232781/
Svetovidov AN, 1941. On the origin of Clupeonella abrau in relation to the development conditions of the herring population in the Caspian, Black and Azov seas. C. R. (Doklady) Acad. Sci. URSS, 31(8):805-809.
Whitehead PJP, 1984. Clupeidae. In: Fishes of the north-eastern Atlantic and the Mediterranean. Vol. 1 [ed. by Whitehead, P. \Bauchot, M. \Hureau, J. \Nielsen, J. \Tortonese, E.]. Paris: UNESCO, 268-281.
Whitehead PJP, 1985. FAO Species Catalogue. Vol. 7. Clupeoid fishes of the world (suborder Clupeioidei). An annotated and illustrated catalogue of the herrings, sardines, pilchards, sprats, shads, anchovies and wolf-herrings. Part 1-Chirocentridae, Clupeidae and Pristigasteridae. FAO Fish. Synop, 125(7/1). 1-303.
Zaitsev YP; Alexandrov BG, 1997. Recent man-made changes in the Black Sea ecosystem. In: Sensitivity to Change: Black Sea, Baltic Sea and North Sea [ed. by Ozsoy, E. \Mikaelyan, A.]., The Netherlands: Kluwer Academic Publishers, 25-31.
OrganizationsTop of page
Lithuania: GTC Nature Research Center, Moksliniku 2, Vilnius, www.gamtostyrimai.lt
ContributorsTop of page
31/05/11 Original text by:
Saulius Stakenas, Gamtos Tyrimu Centro Ekologijos Institutas, Vilnius Lietuva, Lithuania
Reviewers' names are available on request.
Distribution MapsTop of page
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