Hemiculter leucisculus (common sawbelly)

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Adult

Adult female. L = 124 mm. W = 15.7 g. Abs. fecundity = 4600 eggs. Muynak Canal, Low Amu Darya, Uzbekistan, 20th March 2003.

Ernest Khurshut

Identity

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Preferred Scientific Name

Hemiculter leucisculus (Basilewsky, 1855)

Preferred Common Name

common sawbelly

Other Scientific Names

Chanodichthys leucisculus (Basilewsky, 1855)
Culter leucisculus Basilewsky, 1855
Cultriculus akoensis Oshima, 1920
Hemiculter clupeoides Nichols, 1925
Hemiculter eigenmanni (Jordan & Metz, 1913)
Hemiculter kneri Warpachowski, 1887
Hemiculter schrencki Warpachowski, 1888
Hemiculterella eigenmanni Jordan & Mertz, 1913
Kendallia goldsboroughi Evermann & Shaw, 1927
Parapelecus eigenmanni Jordan & Metz, 1913
Squaliobarbus annamiticus Tirant, 1883

International Common Names

English: kili fish; knifefish; Korean sawbelly; minnow; sharpbelly
Chinese: bai tiao; huahua

Local Common Names

Germany: Beilbauch-Weißfish
Iran: kuli-e mordab; shakam tiz; tizeh kuli
Kazakhstan: bileu balyq
Korea, DPR: chili; chiri
Russian Federation/Russian Far East: koreyskaya vostrobryushka; obyknovennaya vostrobryushka
Uzbekistan: oddiy qirraqorin balig'i

Summary of Invasiveness

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Outside of its native area, H. leucisculus was first reported from Central Asia in the Amu Darya Basin in 1958, and in the Syr Darya Basin in 1961. In Iran, it was first found in the Anzali Lagoon (Holcík and Razavi, 1992) where it was probably introduced with Asian carp in 1967.

H. leucisculus is considered as a pest in fish farms where it competes with juveniles of commercial species. It also harms the local fish fauna, and displaces the small, native species, most probably by predation.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Chordata
Subphylum: Vertebrata
Class: Actinopterygii
Order: Cypriniformes
Family: Cyprinidae
Genus: Hemiculter
Species: Hemiculter leucisculus

Description

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The dorsal fin of H. leucisculus has a strong spine. Dorsal spines: 2-3; dorsal soft rays: 7; anal spines: 2-3; anal soft rays: 10-14 (mostly 11-12). Moderate-sized scales that easily fall off. Scale number in the lateral line: 40-59 (mostly 46-52). The gill raker count of the first arch 15-21 (mostly 16-18). Pharyngeal teeth are hooked at the tip: 2,4,4-5,4,2; 2,4,5-4,4,2; 2,3,5-4,3,2. Has a sharp scale-less keel from the throat to the anus. The lateral line goes downward behind the head and then parallels the lower body margin, at the anal fin it curves upward and extends along the middle part of the caudal peduncle. Dark above, green grey and silvery on the back, white on the belly. There is a dark stripe along the upper flank.

Distribution

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The native range of H. leucisculus is East Asia: from Far East Russia and Mongolia in the north, through to China (including Hong Kong and Taiwan) and West Korea, to North Vietnam in the south.

Exotic distribution

H. leucisculus was unintentionally introduced into the Aral Sea Basin, Central Asia. It is currently widespread in the plains of the region up to foothills, and in the drainages of the Amu Darya, Syr Darya, Zarafshan, Qashqadarya, and Tedzhen rivers. In Iran it was reported from the Caspian Sea (Safid River, Aras River, Golestan Province) and Tigris basins. It is thought to be widespread in Iran, although this may be the native fish the Danube bleak (Chalcalburnus chalcoides), which is very similar. H. leucisculus was probably introduced from Central Asia again with Asian carp (Holcík and Razavi, 1992).

Distribution Table

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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.

Last updated: 10 Jan 2020

Continent/Country/Region	Distribution	Origin	First Reported	Invasive	Reference	Notes
Asia
Afghanistan	Present	Introduced			Coad (1981)	Amu Darya Basin (Qonduz province)
China	Present, Widespread	Native			BERG (1949) Bănărescu (1991) Nie et al. (2000)	Yangtze Basin (Jinsha River Basin, Chuanjiang, Minjiang River Basin, Tuojiang River Basin, Jialing River Basin, Wujiang River Basin, Daning River, Xiangxi River), streams draining to Gulf of Chihli (/Bo Hai)
-Anhui	Present	Native			Bănărescu (1991)
-Beijing	Present	Native			Bănărescu (1991)
-Fujian	Present	Native			BERG (1949)
-Guangdong	Present	Native			Bănărescu (1991)
-Guangxi	Present	Native			Bănărescu (1991)
-Guizhou	Present	Native			Bănărescu (1991)
-Hebei	Present	Native			Bănărescu (1991)
-Heilongjiang	Present	Native			BERG (1949) Bănărescu (1991)	Heilong (/Amur) and Songhua (/Sungari) rivers, Xingkai (/Khanka) Lake
-Henan	Present	Native			Bănărescu (1991)
-Hubei	Present	Native			Xie et al. (2001)	Liangzi Lake
-Hunan	Present	Native			Bănărescu (1991)
-Jiangxi	Present	Native			Bănărescu (1991)
-Jilin	Present	Native			Bănărescu (1991)
-Liaoning	Present	Native			BERG (1949) Bănărescu (1991)	Streams draining to Gulf of Chihli (/Bo Hai)
-Shandong	Present	Native			Bănărescu (1991)
-Shanghai	Present	Native			BERG (1949) Bănărescu (1991)
-Yunnan	Present, Widespread	Introduced		Invasive	Junxing (1996)	Most lakes and rivers; First reported: 1963-1965
-Zhejiang	Present	Native			Kottelat (2006)
Hong Kong	Present	Native			BERG (1949) Bănărescu (1991)
Iran	Present, Widespread	Introduced	1967	Invasive	Coad (1995) Abbasi (2005) Patimar and Kiabi (2005) Patimar et al. (2008) CABI (Undated)	Southeastern Caspian Sea Basin, Safid River, Aras River, Golestan Province, Tigris Drainage
Iraq	Present	Introduced			Garstecki and Amr (2011)	Tigris Drainage
Kazakhstan	Present, Widespread	Introduced	1958	Invasive	Mitrofanov (1967) Mitrofanov et al. (1992) Savvaitova and Petr (1999) Mamilov (2011)	First introduced into Almaty Fish farm. Syr Darya, Chu, Ili, and Talas drainages, Balkhash Lake
Mongolia	Present, Localized	Native			BERG (1949) Dulmaa (1999)	Buyr [Nuur] Lake, low Khalkhyn Gol (/Khalkha River)
North Korea	Present	Native			Nikolski (1956)	West Korea
South Korea	Present, Widespread	Native			Nah (1989) Jang et al. (2002) Jang et al. (2003) Junkil et al. (2006)	West Korea, Hwajinpo and Youngsan lakes, Hoedong Reservoir, Kum River
Taiwan	Present	Native			BERG (1949) Bănărescu (1991)
Turkmenistan	Present, Widespread	Introduced		Invasive	Aliev et al. (1963) Shakirova and Nikolaev (1991) Sal'nikov (1998)	First introduced into Karametniyaz fish farm (55 km W from Kerki, Amu Darya River) and the Karakum (/Qaraqum/Garagum) Canal, Murghab River; First reported: 1958-1961
Uzbekistan	Present, Widespread	Introduced	1961	Invasive	Kamilov and Borisova (1966) Borisova (1971) Borisova (1972) Kamilov and Urchinov ZhU (1995) Khurshut (2002) Khurshut and Rakhmatullaeva (2005)	First introduced into Balykchi fish farm, Tashkent Region (60 km SW From Tashkent. Syr Darya and Chirchik rivers junction). Amu Darya, Syr Darya, Zarafshan, Qashqa Darya drainages. Most water bodies in plains of the Aral Sea Basin
Vietnam	Present	Native			CABI (Undated b)	[Ho] Tay Lake, Red River (/Song Koi), Hué River
Europe
Russia	Present				CABI (Undated a)	Present based on regional distribution.
-Russian Far East	Present, Localized	Native		Invasive	BERG (1949) Nikolski (1956) Reshetnikov (1998) Barabanshchikov and Magomedov (2002) Kolpakov et al. (2010)	Native in Amur drainage and Khanka Lake; introduced and invasive in the Artemovka River (Ussuri Bay, Sea of Japan / East Sea) and Razdolnaya River (Peter the Great Bay, Sea of Japan/East Sea)

History of Introduction and Spread

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In 1958, H. leucisculus was unintentionally introduced in Turkmenistan to the Karametniyaz fish farm and the Karakum Canal from the Yangtze River. Then in 1961 it was introduced (again with larvae of Asian carp) in Uzbekistan to the Akkurgan (later Balykchi) fish farm in the Tashkent region. It was transferred, with fish seed, to other fish farms of the region. H. leucisculus has escaped from fish farms and via the extensive network of irrigation canals has spread to the plains of the Aral Sea Basin to the drainages of the Amu Darya, Syr Darya, Zarafshan, Qashqa Darya, and Tedzhen rivers.

In Iran, H. leucisculus was first reported from the Anzali Lagoon by Holcík and Razavi (1992) where it was probably introduced by accident along with Asian carp from Central Asia in 1967 (Holcík and Razavi, 1992). Esmaeili and Gholamifard (2011) reported a significant range extension and the presence in two new drainage basis in Iran, over 370 km southwest of the nearest previous records from the Caspian Sea basin. They also warn that the presence of the species in the Tedzhen River of Turkmenistan may eventually lead to H. leucisculus being found throughout the entire Tedzhen (=Hari) River basin of Iran.

In the Razdolnaya River (Far East Russia) H. leucisculus was unintentionally introduced from Khanka Lake and probably from China as a result of aquaculture activity (Kolpakov et al., 2010). In the Artemovka River (Far East Russia) H. leucisculus was unintentionally introduced from Khanka Lake as a result of aquaculture activity.

Introductions

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Introduced to	Introduced from	Year	Reason	Introduced by	Established in wild through		References	Notes
Introduced to	Introduced from	Year	Reason	Introduced by	Natural reproduction	Continuous restocking	References	Notes
Iran	Former USSR	1967	Stocking (pathway cause)		Yes	No	Holcík and Olah (1992)
Kazakhstan	China	1958-1959	Stocking (pathway cause)		Yes	No	Mitrofanov et al. (1992)	To Almaty fish farm from the Amur Drainage
Turkmenistan	China	1958-1961	Stocking (pathway cause)		Yes	No	Sal'nikov (1998)	Karametniyaz fish farm (55 km W from Kerki, Amu Darya River)
Uzbekistan	China	1961	Stocking (pathway cause)		Yes	No	Borisova (1972); Kamilov and Borisova (1966); Khurshut (2002)	Balykchi fish farm, Tashkent region (60 km SW from Tashkent. Syr Darya and Chirchik rivers junction)

Risk of Introduction

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The main pathway of introduction is the transfer of fish seed between fish farms. H. leucisculus then escapes from fishponds and spreads to the wild via irrigation canals. To help prevent spread of this species all transported fish seed should be properly checked for the presence of H. leucisculus before release into ponds.

Habitat

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H. leucisculus is found in rivers, lakes, reservoirs, canals, and marshes. It keeps to the water surface in stagnant waters.

Habitat List

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Category	Habitat	Presence	Status
Freshwater	Irrigation channels	Secondary/tolerated habitat	Harmful (pest or invasive)
Freshwater	Irrigation channels	Secondary/tolerated habitat	Natural
Freshwater	Lakes	Principal habitat	Harmful (pest or invasive)
Freshwater	Lakes	Principal habitat	Natural
Freshwater	Reservoirs	Principal habitat	Harmful (pest or invasive)
Freshwater	Reservoirs	Principal habitat	Natural
Freshwater	Rivers / streams	Secondary/tolerated habitat	Harmful (pest or invasive)
Freshwater	Rivers / streams	Secondary/tolerated habitat	Natural
Freshwater	Ponds	Principal habitat	Harmful (pest or invasive)
Freshwater	Ponds	Principal habitat	Natural
Brackish	Estuaries	Secondary/tolerated habitat	Natural
Brackish	Lagoons	Secondary/tolerated habitat	Harmful (pest or invasive)
Brackish	Lagoons	Secondary/tolerated habitat	Natural

Biology and Ecology

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Genetics

Molecular studies have been carried out on European cyprinids (Zardoya and Doadrio, 1999) and Asian cyprinids (Liu and Chen, 2003); both papers make reference to H. leucisculus.

The complete mitogenome of H. leucisculus was determined to be 16.62 kbp (Dong, et al., 2015; Xiang et al., 2016).

Reproductive Biology

In Uzbekistan sexual maturity is attained at the age of 2-3 years and at a length of 7-10 cm. Spawning is fractional and takes place from May to July. The average absolute fecundity is 19.9±2.2 thousand eggs and relative fecundity is 583±42 eggs per gram of total body weight (Khurshut, 2013). The average egg diameter is 0.58±0.01 mm. In Kazakhstan the absolute fecundity is 6000-13400 eggs (Mamilov, 2011).

In Erhai Lake, Yunnan Province, China the average batch fecundity is 11934±5921 eggs, while relative batch fecundity is 560±137 eggs g⁻¹ wet weight (Wang et al., 2014).

Physiology and Phenology

Longevity - 7 years.

Nutrition

H. leucisculus is an omnivorous species with a broad feeding plasticity.

In the native area, e.g. in the Amur drainage, H. leucisculus mainly feeds on macrophytes, zooplankton, algae and insects (Markovtsev, 1980). Nikolski (1956) indicated that the basis of diet for this species in Lake Khanka was zooplankton and in particular cladocerans.

The diet of introduced H. leucisculus in Peter the Great Bay (Far East Russia) is discussed in Dolganova et al. (2008).

In water bodies of Kazakhstan and Turkmenistan the diet of this fish consists mostly of vegetative food both by the frequency of encounters and volume. Of the animal food, a significant part consists of crustaceans, insects and their larvae. Cases of predation have also been reported (Borisova, 1971; Shakirova and Nikolaev, 1991; Mitrofanov et al., 1992).

The intestines of H. leucisculus from Uzbekistan mainly contained detritus, and remnants of higher plants and algae (Khurshut and Rakhmatullaeva, 2005). Of the animals, the most frequently recorded were different stages of insects and crustaceans. Of the insects H. leucisculus prefers the larvae of dragonflies, chironomids and the larvae of other dipterans; of crustaceans, cladocerans are reported most frequently.

Natural Food Sources (reported for adults)	Contribution to total food intake (%)
Natural Food Sources (reported for adults)	Contribution to total food intake (%)
Khanka Lake, Russia
Macrophytes	35.2
Terrestrial insects	6.6
Melosira	8.8
Algae	1.7
Chironomids	6.2
Cladocera	40.7
Detritus	0.6
Crustaceans	0.2
Kapchagay reservoir, Kazakhstan
Macrophytes	28.7
Terrestrial insects	24.1
Mysida	17.2
Algae	11.4
Chironomids	9.0
Cladocera	4.8
Copepoda	4.3
Amphipods	0.8
Fish	0.6
Syr Darya River, Uzbekistan
Filamentous algae	27.2
Chironomids	53.2
Hemipterans	3.0
Crustaceans	0.6
Dragonfly larvae	16.0

Climate

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Climate	Status	Description
BS - Steppe climate	Tolerated	> 430mm and < 860mm annual precipitation
BW - Desert climate	Tolerated	< 430mm annual precipitation
Cf - Warm temperate climate, wet all year	Preferred	Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cw - Warm temperate climate with dry winter	Preferred	Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Ds - Continental climate with dry summer	Tolerated	Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter	Preferred	Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)	Latitude South (°S)	Altitude Lower (m)	Altitude Upper (m)
53-20

Air Temperature

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Parameter	Lower limit	Upper limit
Absolute minimum temperature (ºC)		-40
Mean maximum temperature of hottest month (ºC)	12	30
Mean minimum temperature of coldest month (ºC)	-25	14

Water Tolerances

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Parameter	Typical Value	Status	Notes
Depth (m b.s.l.)	0-10	Optimum
Hardness (mg/l of Calcium Carbonate)	15	Optimum	(Mitrofanov et al., 1992)
Salinity (part per thousand)		Optimum	9 tolerated (Kolpakov et al., 2010)
Velocity (cm/h)	0	Optimum	360,000 tolerated
Water pH (pH)	7	Optimum	(Baensch and Fischer, 1998)
Water temperature (ºC temperature)	18-22	Optimum	(Baensch and Fischer, 1998)

Natural enemies

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Natural enemy	Type	Specificity
Ancyrocephalus perplexus	Parasite	not specific
Aspius aspius	Predator	not specific
Bothriocephalus acheilognathi	Parasite	not specific
Bothriocephalus opsariichthydis	Parasite	not specific
Camallanus cotti	Parasite	not specific
Carassotrema koreanum	Parasite	not specific
Carassotrema lamellorchis	Parasite	not specific
Carassotrema megapharyngus	Parasite	not specific
Carassotrema schistorchis	Parasite	not specific
Carassotrema wui	Parasite	not specific
Channa argus argus	Predator	not specific
Channa micropeltes	Predator	not specific
Cleidodiscus brachus	Parasite	not specific
Clonorchis sinensis	Parasite	not specific
Cucullanus cyprini	Parasite	not specific
Dactylogyrus alatoideus	Parasite	not specific
Dactylogyrus anchoratus	Parasite	not specific
Dactylogyrus brachius	Parasite	not specific
Dactylogyrus chenminjungae	Parasite	not specific
Dactylogyrus chini	Parasite	not specific
Dactylogyrus clavaeformis	Parasite	not specific
Dactylogyrus eigenmanni	Parasite	not specific
Dactylogyrus flagellicirrus	Parasite	not specific
Dactylogyrus foliicirrus	Parasite	not specific
Dactylogyrus hemiculteris	Parasite	not specific
Dactylogyrus latituba	Parasite	not specific
Dactylogyrus leucisculus	Parasite	not specific
Dactylogyrus liangshiyani	Parasite	not specific
Dactylogyrus magnihamatus	Parasite	not specific
Dactylogyrus montschadskyi	Parasite	not specific
Dactylogyrus nikolskyi	Parasite	not specific
Dactylogyrus ornithopodus	Parasite	not specific
Dactylogyrus pannosus	Parasite	not specific
Dactylogyrus parabramis	Parasite	not specific
Dactylogyrus paracurvituboides	Parasite	not specific
Dactylogyrus peculiaris	Parasite	not specific
Dactylogyrus proprius	Parasite	not specific
Dactylogyrus pusillus	Parasite	not specific
Dactylogyrus tridigitatus	Parasite	not specific
Dendronucleata petruschewskii	Parasite	not specific
Diplostomum hupehensis	Parasite	not specific
Diplostomum niedashui	Parasite	not specific
Diplozoon hemiculteri	Parasite	not specific
Dollfustrema vaneyi	Parasite	not specific
Echinochasmus beleocephalus	Parasite	not specific
Echinochasmus japonicus	Parasite	not specific
Echinostoma robustum	Parasite	not specific
Gyrodactylus hemiculteris	Parasite	not specific
Haplorchis pumilio	Parasite	not specific
Hebesoma violenteum	Parasite	not specific
Metagonimus yokogawai	Parasite	not specific
Micracanthorhynchina motomurai	Parasite	not specific
Paradiplozoon doi	Parasite	not specific
Paradiplozoon hemiculteri	Parasite	not specific
Parambassis siamensis	Predator	not specific
Procamallanus fulvidraconis	Parasite	not specific
Pseudocapillaria tomentosa	Parasite	not specific
Raphidascaris acus	Parasite	not specific
Rhabdochona hakyi	Parasite	not specific
Rhabdochona jiangxiensis	Parasite	not specific
Rhabdochona longispicula	Parasite	not specific
Sander lucioperca	Predator	not specific
Silurus glanis	Predator	not specific
Triaenophorus amurensis	Parasite	not specific

Notes on Natural Enemies

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In Central Asia, H. leucisculus is predated upon by Sander lucioperca, Silurus glanis and Aspius aspius. However, it is more resistant to predators than similar native species such as Alburnoides oblongus. The last strong spine of the dorsal fin serves as a deterrent to predators.

In Taiwan, H. leucisculus is eaten by the invasive fish species Parambassis siamensis and Channa micropeltes.

Means of Movement and Dispersal

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Accidental Introduction

H. leucisculus has been accidentally transferred with juveniles of Asian carp.

Pathway Causes

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Cause	Notes	Long Distance	Local	References
Stocking	Fries and fingerlings contaminated in stocks of Asian carps	Yes	Yes

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Aquaculture stock	Fries and fingerlings contaminated in stocks of Asian carps	Yes	Yes

Impact Summary

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Category	Impact
Economic/livelihood	Negative
Environment (generally)	Negative

Economic Impact

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H. leucisculus is considered as a pest on fish farms and competitor of juveniles of commercial species. It has also been reported to feed on farmed fish eggs and fry.

Environmental Impact

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Impact on Biodiversity

H. leucisculus may compete with the native species and might also feed on their eggs and fry. It has displaced native species in the Aral Sea Basin due to its higher resistance to predators, the high fecundity and its ability to eat a wide range of food.

Threatened Species

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Threatened Species	Conservation Status	Where Threatened	Mechanism	References
Alburnoides oblongus (Tashkent riffle bleak)	National list(s)	Kazakhstan; Uzbekistan	Competition; Rapid growth	Uzbek Academy of Sciences (2006)
Capoetobrama kuschakewitschi	National list(s)	Kazakhstan; Uzbekistan	Competition; Rapid growth	Uzbek Academy of Sciences (2006)
Leuciscus idus (ide)	No Details		Competition; Rapid growth	Uzbek Academy of Sciences (2006)

Risk and Impact Factors

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Invasiveness

Invasive in its native range
Proved invasive outside its native range
Has a broad native range
Abundant in its native range
Highly adaptable to different environments
Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
Capable of securing and ingesting a wide range of food
Benefits from human association (i.e. it is a human commensal)
Has high reproductive potential
Gregarious

Impact outcomes

Damaged ecosystem services
Negatively impacts aquaculture/fisheries
Reduced native biodiversity
Threat to/ loss of endangered species
Threat to/ loss of native species

Impact mechanisms

Competition (unspecified)
Predation
Rapid growth

Likelihood of entry/control

Difficult to identify/detect in the field
Difficult/costly to control

Uses

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Economic Value

In the native range has a minor commercial importance as a food fish (because it is very bony) and as bait.

In Central Asia and Iran H. leucisculus may be confused with the valuable commercial native species Chalcalburnus chalcoides.

Uses List

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Animal feed, fodder, forage

Bait/attractant

General

Sport (hunting, shooting, fishing, racing)

Human food and beverage

Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Detection and Inspection

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H. leucisculus is easily distinguishable from the native species by the strong spine in the dorsal fin, the elongate anal fin and the curved lateral line, which goes downward behind the head and then parallels the lower body margin, at the anal fin it curves upward and extends along the middle part of the caudal peduncle.

Similarities to Other Species/Conditions

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H. leucisculus may be confused with the native species Alburnoides oblongus, Chalcalburnus chalcoides, and fingerlings of Aspius aspius. However, it is easy to recognize with its strong dorsal spine, an elongate anal fin and lateral line, which goes downward behind the head and then parallels the lower body margin, at the anal fin it curves upward and extends along the middle part of the caudal peduncle.

Prevention and Control

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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

To prevent spread all transported fish seed should be properly checked for the presence of H. leucisculus before release in ponds.

Gaps in Knowledge/Research Needs

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Data on environmental requirements and interactions with native species are insufficient.

References

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Abbasi K, 2005. Studying alien fishes and macrocrustaceans distribution and their effects on rivers and wetlands of the Iranian basin of Caspian Sea. In: II International Symposium "Invasion of Alien species in Holarctic (Borok-2)", 27 September-1 October 2005, Borok, Russia. 194-195

Aliev DS, Verigina IA, Svetovidova AA, 1963. The species composition of fish imported along with grass carp and silver carp from China. In: Problems of fishing of herbivorous fish in the reservoirs of the USSR. Ashgabat, Turkmenistan 178-180

Baensch HA, Fischer GW, 1998. Aquarien Atlas., Germany: Mergus

Banarescu, P., 1991. Zoogeography of fresh waters. Volume 2: distribution and dispersal of freshwater animals in North America and Eurasia, Wiesbaden, Germany: AULA-Verlag.519-1091.

Barabanshchikov EI, Magomedov RA, 2002. Species composition and some features of biology of fishes in the estuaries of Southern Primorye's rivers. Izv. TINRO, 131:179-200

Berg LS, 1949. Freshwater fishes of the U.S.S.R. and adjacent countries. Moscow, Russia: USSR Academy of Sciences

Borisova AT, 1971. Materials on distribution and biology of sawbelly (Hemiculter bleeker) in water bodies of Kalgan-Chirchik fish farm. In: The biological basics of fishery in Uzbekistan [ed. by The biological basics of fishery in Uzbekista]. Fan, Tashkent, Uzbekistan 281-289

Borisova AT, 1972. Accidental invaders in water bodies of Uzbekistan. Journal of Ichthyology, 12(1):49-53

Coad BW, 1981. Fishes of Afghanistan, an Annotated Checklist. National Museum of Natural Sciences (Ottawa) Publications in Zoology, 14:1-26

Coad BW, 1995. Freshwater fishes of Iran. Acta Sci. Nat. Acad. Sci. Brno. 29(1):1-64

Dolganova NT, Kolpakov NV, Chuchukalo VI, 2008. Feeding interactions and foraging of juvenile fish and shrimp in the Estuaries of Peter the Great Bay in the summer-fall period. Russian Journal of Marine Biology, 34(7):482-489

Dong F, Tong GX, Kuang YY, Zheng XH, Sun XW, 1015. The complete mitochondrial genome sequence of Hemiculter leucisculus. Mitochondrial DNA, 26(5), 747-748.

Dulmaa A, 1999. Fish and fisheries in Mongolia. Fish and fisheries at higher altitudes: Asia. [ed. by Petr T]. Rome, Italy: FAO, 187-236. FAO Fisheries Technical Paper No. 385. http://www.fao.org/docrep/003/x2614e/x2614e00.htm

Esmaeili HR, Gholamifard A, 2011. Range extension and translocation for Hemiculter leucisculus (Basilewsky, 1855) (Cyprinidae) in western and northwestern Iran. Journal of Applied Ichthyology, 27(6), 1394-1395.

Esmaeili HR, Gholamifard 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

Garstecki T, Amr Z, 2011. Biodiversity and Ecosystem Management in the Iraqi Marshlands - Screening Study on Potential World Heritage Nomination. Amman, Jordan: IUCN, 189 pp

Holcík J, Olah J, 1992. Fish, fisheries and water quality in Anzali Lagoon and its watershed. Fish, fisheries and water quality in Anzali Lagoon and its watershed. Rome, Italy: FAO. http://www.fao.org/docrep/006/AD192E/AD192E00.htm

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

Jang MH, Joo GJ, Choi SS, 2003. The impact of monsoon on the distribution of fish in a small stream, Korea. Ecohydrology & Hydrobiology, 3(1):87-95

Jang MH, Kyong H, Joo GJ, 2002. Interaction between cyanobacterial bloom and fishes in a small reservoir: laboratory and enclosure experiment. In: International Conference on Limnology of Shallow Lakes. Balaton fured, Hungary: Veszprem

Junkil C, Osamu M, Yasushi S, Kaoru F, 2006. Fish fauna of the Hwajinpo Lake, Korea. Acta Hydrobiologica Sinica, 30(6):633-637

Junxing Y, 1996. The Alien and Indigenous Fishes of Yunnan: A Study on Impact Ways, Degrees and Relevant Issues. In: Conserving China's Biodiversity (II) [ed. by Schei, P. J. \Sung, W. \Yan, X.]. Beijing, China: China Environmental Science Press, 157-168

Kamilov G, Urchinov Zh U, 1995. Fish and fisheries in Uzbekistan under the impact of irrigated agriculture. In: Petr T, ed. Inland fisheries under the impact of irrigated agriculture: Central Asia. FAO Fisheries Circular No. 894, 10-41

Kamilov GC, Borisova AT, 1966. Low-value and pest fish species in "Kalgan-Chirchik" fish farm. In: Vertebrates of Central Asia. Tashkent, Uzbekistan 31-32

Khurshut EE, 2002. Invasive fish species in the Charvak reservoir. In: Biodiversity of the West Tien Shan: protection and efficient use. Tashkent, Uzbekistan: Chinor ENK, 253-257

Khurshut EE, Rakhmatullaeva GM, 2005. Korean sawbelly, Hemiculter leucisculus, a competitor of indigenous species in Uzbekistan. In: II International Symposium "Invasion of Alien species in Holarctic (Borok-2)", 27 September-1 October 2005, Borok, Russia. 198-199

Kolpakov NV, Barabanshchikov EI, Chepurnoi AYu, 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(2):87-94. http://www.maik.ru/abstract/bioinv/10/bioinv0087_abstract.pdf

Kottelat M, 2006. Fishes of Mongolia. A check-list of the fishes known to occur in Mongolia with comments on systematics and nomenclature. Washington DC, USA: El Banc Mundial, 103 pp

Kurshut EE, 2013. Reproductive capacity of Hemiculter leucisculus (Basilewsky, 1855) in Uzbekistan. Communications in Agricultural and Applied Biological Sciences, 78(4), 207-207.

Liu H, Chen Y, 2003. Phylogeny of the east Asian cyprinids inferred from sequences of the mitochondrial DNA control region. Can. J. Zool, 81(12):1938-1946

Mamilov Sh N, 2011. Modern diversity of alien fish species in Chu and Talas river basins. Russian Journal of Biological Invasions, 4(1):65-76

Markovtsev VG, 1980. Nutrition of the Korean sawbelly Hemiculter eigenmanni (Jordan et Metz) in Lake Khanka. Journal of Ichthyology, 20:168-170

Mitrofanov VP, 1967. Characteristics of fish fauna of water bodies of the inundation area of Kapchagai reservoir. In: Biological Basis of Fisheries in Central Asian republics and Kazakhstan. Balkhash, Kazakhstan 193-195

Mitrofanov VP, Dukravets GM, Sidorova AF, 1992. Fishes of Kazakhstan. Vol. 5. Alma-Ata, Kazakhstan: Gylym

Nah CS, 1989. A Comparative Study on Limno-biological Aspects of the Dammed Lakes in the Youngsan River in Korea - Centering on Fish Fauna. The Korean Journal of Ecology, 12(1):51-65

Nie P, Wang GT, Yao WJ, Zhang YA, Gao Q, 2000. Occurrence of Bothriocephalus acheilognathi in cyprinid fish from three lakes in the flood plain of the Yangtze River, China. Diseases of Aquatic Organisms, 41(1):81-82

Nikolski GV, 1956. Fishes of the River Amur basin. Results of the 1945-1949 Amur ichthyological expedition. Izdatelstvo Akad. Nauk SSSR, 551 pp

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

Patimar R, Kiabi BH, 2005. Alien fish species, implications for conservation and management programes; a case study of wetlands of Golestan Province (north of Iran). In: II International Symposium "Invasion of Alien species in Holarctic (Borok-2)", 27 September-1 October 2005, Borok, Russia. 206

Reshetnikov YuS, 1998. Annotated catalog of cyclostomes and fishes of continental waters of Russia. Moscow, Russia: Nauka, 220 pp

Sal'nikov VB, 1998. Anthropogenic Migration of Fish in Turkmenistan. Journal of Ichthyology, 38(8):591-602

Savvaitova KA, Petr T, 1999. Fish and fisheries in Lake Issyk-kul (Tien Shan), River Chu and Pamir lakes. In: T. Petr (ed.) Fish and fisheries at higher altitudes: Asia. FAO Fish. Tech. Pap. No. 385. FAO, Rome, 168-186

Shakirova FM, Nikolaev AA, 1991. Peculiarities of the biology of sawbelly (Hemiculter leucisculus (Basilewsky) in the Murgab basin. Izv. AN TSSR. Biological Sciences Series, 1:24-33

Shed'ko SV, 2001. List of cyclostomes and fishes of fresh waters of Primorye coast. In: Chteniya pamyati Vladimira Yakovlevicha Levanidova Vol. 1., Russia: Dalnauka Publ., 220-249

Uzbek Academy of Sciences, 2006. The Red Data Book of the Republic of Uzbekistan. Volume 2: Animals. Tashkent, Uzbekistan

Wang Teng, Huang Dan, Shen JianZhong, Chen YuShun, Sun GuangWen, Wang HaiSheng, 2014. Batch fecundity and spawning frequency of invasive Hemiculter leucisculus (Basilewsky, 1855) in Erhai Lake, China. Environmental Biology of Fishes, 97(10), 1161-1168. http://rd.springer.com/journal/10641 doi: 10.1007/s10641-013-0205-8

Wang XingLu, Xiang JianGuo, Liu JiaShou, Liu Ming, Wu Lang, Murphy, B. R., Xie SongGuang, 2013. Reduced growth and reproductive investment of Hemiculter leucisculus (Cyprinidae) in a reservoir with introduced icefish Neosalanx taihuensis (Salangidae). Environmental Biology of Fishes, 96(7), 895-903. http://rd.springer.com/journal/10641 doi: 10.1007/s10641-012-0085-3

Xiang D, Ai WM, Peng X, Qiu JB, Chen SB, 2016. Complete mitogenome of Hemiculter leucisculus (Cyprinidae: Cultrinae). Mitochondrial DNA, 27(1), 145-146.

Xie S, Cui Y, Li Z, 2001. Small fish communities in two regions of the Liangzi Lake, China, with or without submersed macrophytes. Journal of Applied Ichthyology, 17(2):89-92

Zardoya R, Doadrio I, 1999. Molecular evidence on the evolutionary and biogeographical patterns of European cyprinids. Journal of Molecular Evolution, 49(2):227-237

Zareian H, Esmaeili HR, Rejad RZ, Vatandoust S, 2015. Hemiculter leucisculus (Basilewsky, 1855) and Alburnus caeruleus Heckel, 1843: new data on their distributions in Iran. Caspian Journal of Environmental Sciences, 13(1), 11-20.

Distribution References

Abbasi K, 2005. Studying alien fishes and macrocrustaceans distribution and their effects on rivers and wetlands of the Iranian basin of Caspian Sea. In: II International Symposium "Invasion of Alien species in Holarctic (Borok-2)", 27 September-1 October 2005, Borok, Russia, 194-195.

Aliev DS, Verigina IA, Svetovidova AA, 1963. The species composition of fish imported along with grass carp and silver carp from China. In: Problems of fishing of herbivorous fish in the reservoirs of the USSR, Ashgabat, Turkmenistan. 178-180.

Bănărescu P, 1991. Zoogeography of fresh waters. Volume 2: distribution and dispersal of freshwater animals in North America and Eurasia. Wiesbaden, Germany: AULA-Verlag. 519-1091.

Barabanshchikov EI, Magomedov RA, 2002. Species composition and some features of biology of fishes in the estuaries of Southern Primorye's rivers. In: Izv. TINRO, 131 179-200.

BERG T, 1949. Swedish Sugar Company's investigations of cultivation technique. 2.e. Crop rotation. (Sockerbolagets odlingstekniska undersökningar. 2.e. Skiftesbruk.). Socker. 107-117.

Borisova AT, 1971. Materials on distribution and biology of sawbelly (Hemiculter bleeker) in water bodies of Kalgan-Chirchik fish farm. In: The biological basics of fishery in Uzbekistan, [ed. by The biological basics of fishery in Uzbekista]. Fan Tashkent, Uzbekistan: 281-289.

Borisova AT, 1972. Accidental invaders in water bodies of Uzbekistan. In: Journal of Ichthyology, 12 (1) 49-53.

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

Coad B W, 1995. Freshwater fishes of Iran. Acta Sci. Nat. Acad. Sci. Brno. 29 (1), 1-64.

Coad BW, 1981. Fishes of Afghanistan, an Annotated Checklist. In: National Museum of Natural Sciences (Ottawa) Publications in Zoology, 14 1-26.

Dulmaa A, 1999. Fish and fisheries in Mongolia. In: FAO Fisheries Technical Paper. [ed. by Petr T]. Rome, Italy: FAO. 187-236. http://www.fao.org/docrep/003/x2614e/x2614e00.htm

Garstecki T, Amr Z, 2011. Biodiversity and Ecosystem Management in the Iraqi Marshlands - Screening Study on Potential World Heritage Nomination., Amman, Jordan: IUCN. 189 pp.

Jang MH, Joo GJ, Choi SS, 2003. The impact of monsoon on the distribution of fish in a small stream, Korea., 3 (1) 87-95.

Jang MH, Kyong H, Joo GJ, 2002. Interaction between cyanobacterial bloom and fishes in a small reservoir: laboratory and enclosure experiment. In: International Conference on Limnology of Shallow Lakes, Balaton fured, Hungary: Veszprem.

Junkil C, Osamu M, Yasushi S, Kaoru F, 2006. Fish fauna of the Hwajinpo Lake, Korea. In: Acta Hydrobiologica Sinica, 30 (6) 633-637.

Junxing Y, 1996. The Alien and Indigenous Fishes of Yunnan: A Study on Impact Ways, Degrees and Relevant Issues. In: Conserving China's Biodiversity (II), [ed. by Schei PJ, Sung W, Yan X]. Beijing, China: China Environmental Science Press. 157-168.

Kamilov G, Urchinov ZhU, 1995. Fish and fisheries in Uzbekistan under the impact of irrigated agriculture. In: Inland fisheries under the impact of irrigated agriculture: Central Asia, [ed. by Petr T]. 10-41.

Kamilov GC, Borisova AT, 1966. Low-value and pest fish species in "Kalgan-Chirchik" fish farm. In: Vertebrates of Central Asia, Tashkent, 31-32.

Khurshut EE, 2002. Invasive fish species in the Charvak reservoir. In: Biodiversity of the West Tien Shan: protection and efficient use, Tashkent, Uzbekistan: Chinor ENK. 253-257.

Khurshut EE, Rakhmatullaeva GM, 2005. Korean sawbelly, Hemiculter leucisculus, a competitor of indigenous species in Uzbekistan. In: II International Symposium "Invasion of Alien species in Holarctic (Borok-2)", 27 September-1 October 2005, Borok, Russia, 198-199.

Kolpakov N V, Barabanshchikov E I, Chepurnoi A Yu, 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 (2), 87-94. http://www.maik.ru/abstract/bioinv/10/bioinv0087_abstract.pdf DOI:10.1134/S2075111710020062

Kottelat M, 2006. Fishes of Mongolia. A check-list of the fishes known to occur in Mongolia with comments on systematics and nomenclature., Washington DC, USA: El Banc Mundial. 103 pp.

Mamilov N Sh, 2011. Modern diversity of alien fish species in the Chu and Talas river basins. Russian Journal of Biological Invasions. 2 (2/3), 112-119. DOI:10.1134/S207511171102007X

Mitrofanov VP, 1967. Characteristics of fish fauna of water bodies of the inundation area of Kapchagai reservoir. In: Biological Basis of Fisheries in Central Asian republics and Kazakhstan, Kazakhstan, 193-195. http://Balkhash

Mitrofanov VP, Dukravets GM, Sidorova AF, 1992. Fishes of Kazakhstan., 5 Alma-Ata, Kazakhstan: Gylym.

Nah CS, 1989. A Comparative Study on Limno-biological Aspects of the Dammed Lakes in the Youngsan River in Korea - Centering on Fish Fauna. In: The Korean Journal of Ecology, 12 (1) 51-65.

Nie P, Wang G T, Yao W J, Zhang Y A, Gao Q, 2000. Occurrence of Bothriocephalus acheilognathi in cyprinid fish from three lakes in the flood plain of the Yangtze River, China. Diseases of Aquatic Organisms. 41 (1), 81-82. DOI:10.3354/dao041081

Nikolski GV, 1956. Fishes of the River Amur basin. In: Results of the 1945-1949 Amur ichthyological expedition, Izdatelstvo Akad Nauk SSSR. 551 pp.

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

Patimar R, Kiabi BH, 2005. Alien fish species, implications for conservation and management programes; a case study of wetlands of Golestan Province (north of Iran). In: II International Symposium "Invasion of Alien species in Holarctic (Borok-2)", 27 September-1 October 2005, Borok, Russia, 206.

Reshetnikov YuS, 1998. Annotated catalog of cyclostomes and fishes of continental waters of Russia., Moscow, Russia: Nauka. 220 pp.

Sal'nikov VB, 1998. Anthropogenic Migration of Fish in Turkmenistan. In: Journal of Ichthyology, 38 (8) 591-602.

Savvaitova KA, Petr T, 1999. Fish and fisheries in Lake Issyk-kul (Tien Shan), River Chu and Pamir lakes. In: Fish and fisheries at higher altitudes: Asia. FAO Fish. Tech. Pap. No. 385, [ed. by Petr T]. Rome, FAO. 168-186.

Shakirova FM, Nikolaev AA, 1991. Peculiarities of the biology of sawbelly (Hemiculter leucisculus (Basilewsky) in the Murgab basin. In: Izv. AN TSSR. Biological Sciences Series, 1 24-33.

Xie S, Cui Y, Li Z, 2001. Small fish communities in two regions of the Liangzi Lake, China, with or without submersed macrophytes. Journal of Applied Ichthyology. 17 (2), 89-92. DOI:10.1046/j.1439-0426.2001.00248.x

Links to Websites

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Website	URL	Comment
Fisheries and Aquaculture Department FAO	http://www.fao.org/fishery/introsp/search/en
Fishes of Uzbekistan	http://uznix.narod.ru/sci/fkey/uzfishlst.html
Freshwater Fishes of Iran	http://www.briancoad.com
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.

Contributors

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16/10/2016 Updated by:

Ernest Khurshut, Center of Genomics and Bioinformatics, Uzbek Academy of Sciences, 2 Universitet St., Tashkent 111215. Toshkent vil., Qibray tum., Uzbekistan

25/07/11 Original text by:

Ernest Khurshut, Institute of Zoology, Uzbek Academy of Sciences, 1 Niyazov St., Tashkent 100095. Uzbekistan

Reviewers' names are available on request.

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Hemiculter leucisculus (common sawbelly)

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