Clupeonella cultriventris (Black Sea kilka)
Datasheet Types: Natural enemy, Invasive species
Abstract
This datasheet on Clupeonella cultriventris covers Identity, Overview, Distribution, Dispersal, Diagnosis, Biology & Ecology, Environmental Requirements, Natural Enemies, Impacts, Uses, Prevention/Control, Further Information.
Identity
- 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
- EnglishAzov Sea kilkaCaspian Sea kilkakilkaPonto-Caspian kilka
- Spanishespadín del Mar Caspioespadín del Mar de Azovespadín del Mar Negro
- Frenchclupeonelle
- Russianchernomorsko-kaspiyskaya kilka (tulka)obyknovennaya tyulka, obyknovennaya kilka
- Portugueseespadilha do Cáspioespadilha-do-Mar-Negro
- Local Common Names
- Bulgariatzatza
- Czech Republickilka kaspická
- Denmarkkaspisk sild
- GermanyKaspische Sprotte
- Irankilka-e-maamooli
- Italypapaline del Caspio
- Kazakhstankilkas tyulka lar
- Netherlandszwarte-zeesprot
- Polandkilka kaspijska
- Romaniagingirica
- Swedenkaspisk skarpsill
- Turkeykilka baligi
- Turkmenistanadaty kulke balyk
- Ukrainetyulka
Pictures

Adults
Clupeonella cultriventris (black sea kilka, or black sea sprat); adults. Danube delta, Ukraine. February 2011.
©Yuriy Kvach-Ukraine/via wikipedia - CC BY-SA 3.0

Adults
Clupeonella cultriventris (black sea kilka, or black sea sprat); adults. Danube delta, Ukraine. February 2011.
©Yuriy Kvach-Ukraine/via wikipedia - CC BY-SA 3.0
Summary of Invasiveness
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 Tree
Notes on Taxonomy and Nomenclature
Nordmann (1840), Borodin (1896) and Svetovidov (1941) detail the taxonomy and nomenclature of Clupeonella cultriventris.
Description
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).
Distribution
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 Map
Distribution Table
History of Introduction and Spread
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 Introduction
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).
Means of Movement and Dispersal
Natural Dispersal (Non-Biotic)
Construction of dam cascades in the main rivers of spawning of C. cultriventris have coincided with their dispersal (Leppäkoski et al., 2002; Kiyashko et al., 2006.).
Accidental Introduction
Possible as a contaminant of ballast water, but no specific cases are cited.
Pathway Vectors
Pathway vector | Notes | Long distance | Local | References |
---|---|---|---|---|
Water (pathway vector) | All life stages by natural dispersal | Yes |
Similarities to Other Species/Conditions
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.
Habitat
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 List
Category | Sub category | Habitat | Presence | Status |
---|---|---|---|---|
Brackish | Inland saline areas | Secondary/tolerated habitat | Natural | |
Littoral | Coastal areas | Principal habitat | Natural | |
Freshwater | ||||
Freshwater | Irrigation channels | Secondary/tolerated habitat | Natural | |
Freshwater | Lakes | Secondary/tolerated habitat | Natural | |
Freshwater | Reservoirs | Secondary/tolerated habitat | Natural | |
Freshwater | Rivers / streams | Principal habitat | Natural | |
Brackish | ||||
Brackish | Estuaries | Principal habitat | Natural | |
Brackish | Lagoons | Principal habitat | Natural | |
Marine |
Biology and Ecology
Genetics
The total mitochondrial genome of C. cultriventris has been sequenced (http://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?id=501719).
Reproductive Biology
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
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).
Growth
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).
Nutrition
Feeding type
Heterotrophic (live organic substances).
Feeding behavior
Preying and selected capture of food items.
Food spectrum
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).
Climate
Climate type | Description | Preferred or tolerated | Remarks |
---|---|---|---|
BS - Steppe climate | > 430mm and < 860mm annual precipitation | Tolerated | |
Cf - Warm temperate climate, wet all year | Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year | Preferred | |
Df - Continental climate, wet all year | Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year) | Tolerated |
Latitude/Altitude Ranges
Latitude North (°N) | Latitude South (°S) | Altitude lower (m) | Altitude upper (m) |
---|---|---|---|
36-60 |
Water Tolerances
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 enemy of
Notes on Natural Enemies
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).
Natural enemies
Natural enemy | Type | Life stages | Specificity | References | Biological control in | Biological control on |
---|---|---|---|---|---|---|
Acipenser persicus (persian sturgeon) | Predator | All Stages | not specific | |||
Acipenser stellatus (starry sturgeon) | Predator | All Stages | not specific | |||
Esox lucius (pike) | Predator | All Stages | not specific | |||
Lota lota | Predator | All Stages | not specific | |||
Perca fluviatilis (perch) | Predator | All Stages | not specific | |||
Sander lucioperca (pike-perch) | Predator | All Stages | not specific | |||
Sander volgensis | Predator | All Stages | not specific |
Impact Summary
Category | Impact |
---|---|
Economic/livelihood | Positive and negative |
Environment (generally) | Positive and negative |
Impact
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 Factors
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
Gregarious
Impact outcomes
Altered trophic level
Conflict
Damaged ecosystem services
Impact mechanisms
Competition - monopolizing resources
Competition (unspecified)
Rapid growth
Likelihood of entry/control
Difficult to identify/detect as a commodity contaminant
Difficult/costly to control
Uses
Economic Value
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).
Social Benefit
Salted and tinned C. cultriventris in southern Ukraine and southern Russia are traditional foods, especially as part of Lent diets.
Environmental Services
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 List
General > Research model
General > Sociocultural value
Detection and Inspection
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.
Prevention and Control
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Prevention
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)’.
Rapid response
This is established at the national level.
Control
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).
Movement control
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.
Biological control
There are no known forms of biological control specific to C. cultriventris.
Chemical control
The only effective method of fish eradication is the application of rotenone, a pesticide that is also toxic to non-target native species.
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 Needs
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).
Links to Websites
Name | URL | Comment |
---|---|---|
DAISIE Delivering Alien Invasive Species Inventories for Europe | http://www.europe-aliens.org/index.jsp | |
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway | https://doi.org/10.5061/dryad.m93f6 | Data source for updated system data added to species habitat list. |
Global register of Introduced and Invasive species (GRIIS) | http://griis.org/ | Data source for updated system data added to species habitat list. |
ISSG database global Invasive Species Database) | http://www.issg.org |
Organizations
Name | Address | Country | URL |
---|---|---|---|
GTC Nature Research Center | Moksliniku 2, Vilnius | Lithuania | www.gamtostyrimai.lt |
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