Neogobius melanostomus (round goby)

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Picture

Title

Caption

Copyright

Adult

Neogobius melanostomus (round goby); adult. USA.

©Eric Engbretson/US Fish & Wildlife Service/Bugwood.org - CC BY-NC 3.0 US

Lateral view

Lateral view of a Round goby from the Gulf of Gdansk, Baltic Sea, Poland.

©Michal Grabowski

Head

Close-up of the head of a Round goby.

©Michal Grabowski

Identity

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Preferred Scientific Name

Neogobius melanostomus (Pallas, 1814)

Preferred Common Name

round goby

Other Scientific Names

Apollonia melanostomus (Illin, 1927)
Gobius melanostomus (Pallas, 1811)

International Common Names

English: black spotted goby
Spanish: gobio de boca negra

Local Common Names

Denmark: sortmundet kutling
Germany: Shwarzmund Grundel
Netherlands: zwartbekgrondel
Russian Federation: bychok-krugyak
Sweden: svartmunnad smörbult

Summary of Invasiveness

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N. melanostomus, a small (up to 25 cm length) benthic species of Ponto-Caspian origin, is one of the most successful fish invaders of the recent decades. Since its first recording outside the natural range i.e. in the Moskva River in the 1980s, the species has inhabited several countries in Europe (Poland, Hungary, Slovakia, Austria, Germany, Estonia, the Netherlands) as well as the Great Lakes of North America. The ballast-water transport and possibly as eggs attached to ship hulls and barges are the most probable vectors of N. melanostomus transmission. Other human-mediated factors like alteration of natural freshwater and marine habitats e.g. rip-rap river bank, artificial reefs (wave breakers), industrial harbours, dam reservoirs, seem to promote its establishment in newly invaded areas. N. melanostomus possesses many biological attributes that facilitate its invasion i.e. broad native range, wide tolerance for environmental conditions (euryhaline species), habitat generalist, opportunistic feeder, early maturation, multiple spawning during prolonged breeding season, male care for eggs deposited in shelter, aggressive behaviour. N. melanostomus ability to survive in harsh conditions, also in degraded environments, has helped to increase its competitive advantage compared to native species. The invasion of N. melanostomus resulted in varied alteration of ecosystem processes e.g. the diet shift among predators and changes in food web structure was noted in the Gulf of Gdansk (the Baltic Sea); in the Great Lakes the recruitment failure of some native species and consequently decline of their population coincided with N. melanostomus presence. N. melanostomus is a voracious predator that feeds on variety of invertebrate benthic organisms, fish eggs, but predominantly on molluscs (e.g. zebra and quagga mussel) that are known to accumulate persistent contaminants from the sediment. As many native game and commercial fish species prey on N. melanostomus the greater bioaccumulation of toxins such as PCBs in the food chain is possible and the fish diet of humans is a health concern.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Chordata
Subphylum: Vertebrata
Class: Actinopterygii
Order: Perciformes
Suborder: Gobioidei
Family: Gobiidae
Genus: Neogobius
Species: Neogobius melanostomus

Notes on Taxonomy and Nomenclature

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The subfamily Goibionellinae (Nelson, 1994), was previously labelled Benthophilinae. Based on results of electrophoretic studies, Dobrovolov et al. (1995) suggested a rearrangement of Gobiidae that moved Neogobius melanostomus to Apollonia melanostomus. The specific epithet melanostomus means ‘black-mouthed’ from the Greek melanos, black, and soma, mouth, but might be thought to refer to the conspicuous black spot typically present on the first dorsal fin (Pinchuk et al., 2003).

Description

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N. melanostomus is small, up to 25 cm in total length. Its pelvic fins form a very characteristic suction disc on the ventral surface. The pelvic disc is 0.6-0.8 times the abdomen length. The body is scaled on the parietal region, nape, back, throat, abdomen, pectoral fin peduncles, and one quarter of the gill covers. The head is wide (as or wider than deep) and relatively big (22-23% of body length). Eyes are large and protrude slightly from the top of the head. The angle of the jaw is below the anterior quarter of the eye. The lower jaw is not prominent. The anterior dorsal fin has 5-7 spines, usually 6, and the posterior dorsal fin has one spine and 13-16 soft rays. The anal fin has one spine and 11-14 soft rays. The pectoral fins have 17-20 soft rays. N. melanostomus lacks a gas bladder and chemoreceptors. Neuromasts are present throughout the body and head. N. melanostomus lacks a visible lateral line. Males are larger than females. Both sexes have an erectile urogenital papilla between the anus and the base of the anal fin. Coloration: yellowish-grey, with lateral blotches; first dorsal fin with large black spot in posterior part; breeding males are almost black and their median fins are more elongated and white-edged (Pinchuk et al., 2003).

Distribution

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N. melanostomus is widely distributed throughout the Ponto-Caspian basins, in freshwater, estuarine and coastal habitats. It is found in the Sea of Azov, the Caspian Sea, Black Sea and Sea of Marmara near Istanbul, Turkey. It also inhabits tributaries of Black and Caspian seas, including the Dniester River as far as the Smotrich River (near the city Kamenets-Podolsky, Ukraine); in the Prut River in the city Chernovtsy (Ukraine), the Southern Bug River in the city Lodizhino (Ukraine); in the Dnieper River in the city Dnepropetrovsk (Ukraine); the Don River in the city Rostov (Russia); in the Volga River to the Volgograd Reservoir (lower reach of the river) but since 1968, it established also in the Kuybyshev Reservoir (middle stretch of the river) (Russia); the Ural River to the Sakma River (Pinchuk et al., 2003). The recent upstream expansion of the species has been observed in the Dniestr, Dniepr, Southern Bug, Volga rivers and its tributaries (Smirnov, 1986; Moskalova, 1996). In the Danube River it was found earlier as far as the town of Vidin (Bulgaria), but has recently spread upstream from the Iron Gate (Simonovic et al., 1998) and even farther reaching Vienna (Austria) (Wiesner et al., 2000).

Outside the Ponto-Caspian basins N. melanostomus invaded several locations in southern coast of the Baltic Sea, and has been reported from Poland (Skóra and Stolarski, 1993), Germany (Corkum et al., 2004), Estonia, Finland, Lithuania (Ojaveer, 2006). Since 2004 it is present in the North Sea basin (Norway) (Van Beek, 2006).

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: 17 Dec 2021

Continent/Country/Region	Distribution	Last Reported	Origin	First Reported	Invasive	Reference	Notes
Asia
Azerbaijan	Present, Localized		Native			Pinchuk et al. (2003)	in rivers and lakes south of the country
Georgia	Present, Localized		Native			Pinchuk et al. (2003)	In the Black Sea
Iran	Present, Localized		Native			Pinchuk et al. (2003)
Kazakhstan	Absent, Formerly present	1960				Pinchuk et al. (2003)	Introduced to Aral Sea now reported as extinct
Turkey	Present, Localized		Native			Pinchuk et al. (2003)	in the Bosphorus, Sea of Marmara and some lakes
Turkmenistan	Present, Localized		Native			Pinchuk et al. (2003)	In Caspian Sea
Uzbekistan	Absent, Formerly present	1960				Pinchuk et al. (2003)	introduced to Aral Sea, now reported as extinct
Europe
Austria	Present, Localized	2000	Introduced		Invasive	Wiesner et al. (2000) Seebens et al. (2017)	in the Danube River
Belarus	Present, Localized	1998	Introduced		Invasive	Gulugin and Kunitsky (1999)	in the Dnieper River and Pripyat River
Belgium	Present		Introduced	2010		Seebens et al. (2017)
Bulgaria	Present, Widespread		Native			Pinchuk et al. (2003)
Croatia	Present		Introduced	1990		Seebens et al. (2017)
Czechia	Present		Introduced	2008		Seebens et al. (2017)
Denmark	Present		Introduced	2008		Seebens et al. (2017)
Estonia	Present, Localized	2002	Introduced		Invasive	Ojaveer (2006) Seebens et al. (2017)	NE part of the Gulf of Riga (Baltic Sea)
Federal Republic of Yugoslavia	Present, Localized		Native		Invasive	Simonovic et al. (1998)
Finland	Present, Few occurrences		Introduced		Invasive	Ojaveer (2006) Seebens et al. (2017)	coastal waters of the Baltic Sea
Germany	Present, Few occurrences	1999	Introduced			Corkum et al. (2004) Seebens et al. (2017)	near Rugia Island and northern coast
Hungary	Present, Localized	2001	Introduced		Invasive	Guti et al. (2003)	in the Danube River
Italy	Present		Introduced	2012		Seebens et al. (2017)
Latvia	Present		Introduced	2004		Seebens et al. (2017)
Lithuania	Present, Few occurrences		Introduced		Invasive	Ojaveer (2006) Seebens et al. (2017)	coastal waters of the Baltic Sea
Netherlands	Present, Few occurrences	2004	Introduced		Invasive	Beek (2006) Seebens et al. (2017)	The Lek River
Norway	Present		Introduced	1930		Seebens et al. (2017)
Poland	Present, Localized	1990	Introduced		Invasive	Skóra and Stolarski (1993) Seebens et al. (2017)	coastal waters of the Baltic Sea; lower section of the Vistula River
Romania	Present		Native			Pinchuk et al. (2003)	Black Sea coast and lagoons; the Danube River basin
Russia	Present		Native		Invasive	Pinchuk et al. (2003) Seebens et al. (2017)	expand its previous natural distribution e.g. in the Volga River basin
-Central Russia	Present, Localized	1985	Introduced		Invasive	Pinchuk et al. (2003)	the Moskva River; the Volgograd Reservoir
-Southern Russia	Present, Widespread		Native		Invasive	Pinchuk et al. (2003)	native to the Caspian and Black seas; lower section of Volga and Don rivers
Serbia	Present, Localized	1997	Introduced		Invasive	Simonovic et al. (1998) Seebens et al. (2017)	the Danube River basin
Slovakia	Present, Localized	2003	Introduced		Invasive	CABI (Undated)	the Danube River basin; Original citation: Stránai and , (2004)
Sweden	Present		Introduced	2008		Seebens et al. (2017)
Ukraine	Present		Native		Invasive	Pinchuk et al. (2003) Seebens et al. (2017)	Native to the Black Sea, lower section of the Dneiper River and Dniestr River; expand farther upstream up to Kiev.
North America
Canada	Present					CABI (Undated a) Seebens et al. (2017)	Present based on regional distribution.
-Ontario	Present, Localized	1990	Introduced		Invasive	Jude et al. (1992)	St. Clair River, Detroit River
United States	Present					CABI (Undated a) Seebens et al. (2017)	Present based on regional distribution.
-Illinois	Present, Localized	1993	Introduced		Invasive	Jude (1997)	Calumet River, Lake Michigan
-Indiana	Present	1996	Introduced		Invasive	Jude (1997)	Lake Michigan
-Michigan	Present, Localized	1990	Introduced		Invasive	Jude (1997)	e.g. St Clair River, Lake Erie, Shiawassee River
-Minnesota	Present	1995	Introduced		Invasive	Jude (1997)	Lake Superior
-New York	Present	1995	Introduced		Invasive	Jude (1997)	Lake Ontario
-Ohio	Present, Localized	1993	Introduced		Invasive	Jude (1997)	Lake Erie
-Pennsylvania	Present	1996	Introduced		Invasive	Jude (1997)	Lake Erie

History of Introduction and Spread

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In the Baltic Sea basin N. melanostomus started its invasion probably from the late 1980s, when in 1990 few individuals 3-4 years old have been found for the first time in the Gulf of Gdansk (the Baltic Sea, Poland) (Skóra and Stolarski, 1993).The species soon spread along the Polish coast (Sapota, 2004) but also entered the Vistula River as far as 130km upstream from the mouth (Kostrzewa and Grabowski, 2002). It has been recently reported from several other places in the southern coastal waters of the Baltic Sea (Ojaveer, 2006) and in the North Sea basin (Van Beek, 2006). The most probable route of N. melanostomus migration to the southern part of Baltic Sea is the so-called northern corridor (Bij de Vaate et al., 2002) consisting the Volga River, Rybinsky Reservoir, lakes Ladoga and Onega connected by artificial canals with the Gulf of Finland (Sapota, 2004). Its introduction to the Baltic Sea is usually related to ballast-water transport (Sapota, 2004) and possibly as eggs attached to the hulls of barges. N. melanostomus invasion to the Baltic Sea was preceeded by its previous expansion in the Volga River system e.g. in the late 1980s it was found in the Moscow River basin (tributary of the upper stretch of the Volga) (Pinchuk et al., 2003).

Its presence in North America was first recorded in 1990 in the St Clair River at Sarnia (Ontario) (Jude et al., 1992). Since 1993, N. melanostomus has been noted in western Lake Superior, Lake Michigan, Lake Huron, Lake Erie and Lake Ontario (Charlebois et al., 1997). It is thought that the species was transported to the Great Lakes in North America initially in ballast water of transatlantic ships, which could have come from the Black Sea basin or from the Baltic, and further distributed within the lakes system by similar means (Jude, 1997).

The precise source of the both introductions outside the Ponto-Caspian basin have not been identified yet. Based on results of mitochondrial DNA studies neither the Great Lakes or the Baltic populations originated from the northern Black Sea (Dougherty et al., 996; Dillion andStepien, 2001). The level of genetic variation indicates that founding populations were relatively large.

The expansion along the Danube River farther upstream from the Iron Gate (suggested border of its natural range in the Danube) is also observed (Simonovic et al., 1998; Wiesner et al., 2000; Guti et al., 2003; Stránai and Andreji, 2004). It is supposed that the expansion of N. melanostomus in the Danube may be the result of natural migration encouraged by water transport and human-mediated alteration of river banks e.g. rip-rap habitats and other harbour artificial substrates seem to be preferred by N. melanostomus for shelter (Ahnelt et al., 1998; Wiesner, 2005). This also explains the jump-like dispersal observed in the Danube system.

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
Austria		2000			Yes	No	Wiesner et al. (2000)
Belarus	Ukraine	1998			Yes	No	Gulugin and Kunitsky (1999)
Canada	Eastern Europe	1990	Interbasin transfers (pathway cause)		Yes	No	Jude et al. (1992)
Estonia		2002			Yes	No	Ojaveer (2006)
Finland		2000s?			No	No
Germany		1999			Yes	No	Corkum et al. (2004)
Hungary		2001			Yes	No	Guti et al. (2003)
Lithuania		2000s?			No	No	Ojaveer (2006)
Norway		2004			No	No	Beek GCWvan (2006)
Poland	Russian Federation	1990	Interbasin transfers (pathway cause); Interconnected waterways (pathway cause)		Yes	No	Sapota (2004); Skóra and Stolarski (1993)
Slovakia		2003			Yes	No
USA	Eastern Europe	1990	Interbasin transfers (pathway cause)		Yes	No	Jude et al. (1992)

Risk of Introduction

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The further expansion of N. melanostomus to Western Europe may be expected from the Danube River through system of canals that connect it with the main river (the right tributary of the Rhine River, the North Sea basin). The other route to the west and north of Europe is possiblly along the Baltic and North Sea coasts. N.melanostomus spread may be due to natural migration combined with passive dispersal by ships and barges.

In North America N. melanostomus has already spread into an inland river in Ontario (Running Creek) and down the Chicago Sanitary and Shipping Canal en route to the Mississippi River (Charlebois et al., 2001).

Habitat

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N. melanostomus is a benthic euryhaline species, inhabiting freshwaters of rivers and lakes to brackish polyhaline salinities. In the Black Sea it is found in coastal shallows on stony or sandy bottom and mussel beds at depths of 1.5-20m, being particularly abundant in shallow bays and estuaries coinciding with the distribution of Dreissena sp. (Pinchuk et al., 2003). In the Gulf of Gdansk it inhabits also softer well vegetated substrates, but artificial reefs (wave breakers), piers, rocky and stony areas are the most favourable (Sapota and Skóra, 2005). In the Danube River, N. melanostomus occurred in the highest numbers on gravel-rocky and stony substrates (Jurajda et al., 2005) and in industrial harbours (Wiesner, 2005). In the Great Lakes systemN. melanostomus is typically confined to lakes and connected channels or navigational waterways (Corkum et al., 2004), mostly found on pebbles with aquatic macrophytes from shallows to 7m depth, but also on sandy bottoms (Charlebois et al., 1997).

Habitat List

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Category	Habitat	Presence	Status
Multiple
Littoral	Coastal areas	Principal habitat	Natural
Freshwater
Freshwater	Lakes	Secondary/tolerated habitat	Harmful (pest or invasive)
Freshwater	Lakes	Secondary/tolerated habitat	Natural
Freshwater	Reservoirs	Principal habitat	Harmful (pest or invasive)
Freshwater	Rivers / streams	Principal habitat	Harmful (pest or invasive)
Freshwater	Rivers / streams	Principal habitat	Natural
Brackish
Brackish	Estuaries	Principal habitat	Natural
Brackish	Lagoons	Principal habitat	Natural
Marine
Marine	Inshore marine	Principal habitat	Harmful (pest or invasive)
Marine	Inshore marine	Principal habitat	Natural
Marine	Benthic zone	Principal habitat	Harmful (pest or invasive)
Marine	Benthic zone	Principal habitat	Natural

Biology and Ecology

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Genetics

N. melanostomus possesses 46 acrocentric chromosomes (Pinchuk et al., 2003). The species has a relatively low level of genetic variability (Charlebois et al., 1997; Pinchuk et al., 2003). The genetics of this species have been extensively studied, see Brown and Stepien (2008) for further details.

Reproductive Biology

In the Ponto-Caspian region (in the native population of N. melanostomus) males become mature during the third year of life and females are mature at the age of two. Fecundity ranges from 200 to 5221 eggs depending on female size. The pale yellow to orange pyriform eggs are about 3.9 by 2.8 mm. They are described as oligoplastic and polylecithal, rich in protein and lipid. The breeding season is prolonged and can last from April to September. Females can spawn up to six times during breeding period. Eggs are deposited in a net on the underside of different structures e.g. rocks, logs, roots, in beer cans, empty shells of molluscs etc, and the nest is defended by the male. Several females may deposit eggs in the same site. Sound production features in the courtship and territorial behaviour is reported for both sexes. Size under hatching is 5.5-5.7 mm. Larvae resemble adults at hatching and appear to be benthic, since they have no swim bladder (Pinchuk et al., 2003).

N. melanostomus from new colonized areas (Baltic Sea, Great Lakes) tend to be smaller, mature earlier at a smaller size compared with the populations in native range. Dwarf morphs are observedwhich direct more energy to reproduction than the normal morphs in native populations (Corkum at al., 2004).

Physiology and Phenology

N. melanostomus is eurythermal and euryhaline species. It can withstand very low level of dissolved oxygen. Overall, the species has wide tolerance of several environmental abiotic factors that enhance its invasiveness. The primary diet of N. melanostomus includes molluscs, crustaceans, worms, fish eggs, small fish, and insect larvae (Pinchuk et al., 2003).

Associations

It is often reported as associated with Dreissena sp. both in native and invaded areas. In rocky substrate habitats, greater than 100 mm N. melanostomus feeds almost exclusively on zebra mussel (Dreissena polymorpha). In addition, since the invasion of quagga mussel (Dreissena bugensis), which tolerates deeper and cooler waters, another source of mussels may be available for N. melanostomus in deeper, soft regimented areas of the Great Lakes, where zebra mussels were prevented from colonizing because of lack of hard substrate (Jude, 1997).

Climate

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Climate	Status	Description	Remark
Cs - Warm temperate climate with dry summer	Preferred	Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Ds - Continental climate with dry summer	Tolerated	Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)

Latitude/Altitude Ranges

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Latitude North (°N)	Latitude South (°S)	Altitude Lower (m)	Altitude Upper (m)
40-60

Water Tolerances

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Parameter	Status	Notes
Dissolved oxygen (mg/l)	Optimum	0.3-0.9 tolerated. The threshold value depending on the mass of fish. Tolerate low oxygenation
Salinity (part per thousand)	Optimum	1-40.6 tolerated. A euryhaline species
Velocity (cm/h)	Optimum	Prefers rather stagnant waters or those of a slow flow
Water temperature (ºC temperature)	Optimum	-1-30 tolerated. A eurythermal species

Natural enemies

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Natural enemy	Type	Life stages	Specificity	References
Ambloplites rupestris	Predator	Aquatic\|Adult	not specific	Jude (1997)
Anguillicoloides crassus	Parasite		not specific	Rolbiecki l (2006)
Cryptocotyle concavum	Parasite		not specific	Kvach and Skóra (2007)
Diplostomum spathaceum	Parasite		not specific	Kvach and Skóra (2007)
Hysterothylacium aduncum	Parasite		not specific	Rolbiecki l (2006)
Lota lota	Predator	Aquatic\|Adult	not specific	Jude (1997)
Micropterus dolomieu	Predator	Aquatic\|Adult	not specific	Jude (1997)
Morone chrysops	Predator	Aquatic\|Adult	not specific	Corkum et al. (2004)
Neochasmus umbellus	Parasite	Aquatic\|Larval	not specific	Kvach and Stepien (2008)
Noturus flavus	Predator	Aquatic\|Adult	not specific	Jude (1997)
Perca flavescens	Predator	Aquatic\|Adult	not specific	Jude (1997)
Phalacrocorax carbo	Predator	Aquatic\|Adult	not specific	Bzoma and Stempniewicz (2001)
Sander vitreus	Predator	Aquatic\|Adult	not specific	Corkum et al. (2004)

Notes on Natural Enemies

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In the Great Lakes of North America N. melanostomus is a common prey of small bass (Micropterus dolomieu), rock bass (Ambloplites rupestris,) stonecat (Noturus flavus), burbot (Lota lota) and yellow perch (Perca flavescens), but is a minor component in the diet of walleye (Stizostedion vitreum) and white bass (Morone chrysops) (Jude,1997; Corkum et al., 2004).

Although fish predators on N. melanostomus in the Gulf of Gdansk (Baltic Sea) are unknown, great cormorants (Phalacrocorax carbo) feed predominantly (72%) on N. melanostomus (Bzoma and Stempniewicz, 2001).

Parasites of N. melanostomus in invaded areas i.e. Gulf of Gdansk (the Baltic Sea, Poland) consist of 12 species. The core of the parasite fauna comprises two species: Cryptocotyle concavum and Diplostomum spathaceum; secondary species are absent; satellite species include Cercariae and Ergasilus sieboldi; rare species are Acanthocephalus lucii, Anguillicola crassus, Bothriocephalus sp., Dichelyne minutus, Hysterothylacium aduncum, Pomphorhynchus laevis, Piscicola geometra, and Tylodelphys clavata. Fifty percent of metazoa parasites that occurred in the invasive N. melanostomus in the Gulf of Gdansk are also typically found in the native Gulf of Gdansk gobiids. N. melanostomus hosts common fish parasite species: C. concavum and D. minutus, but none that are unique to the species and no Ponto-Caspian parasites (Kvach and Skóra, 2007). In the Vistula Lagoon (Poland) parasitological examination revealed the presence of 20 parasitic taxa. The most frequent were nematodes Hysterothylacium aduncum and Anguillicola crassus (Rolbiecki, 2006). In North America, 76% of round gobies were infected with one or more parasites, mostly freshwater. Diplostomum spathacum a species typical of freshwater in the Ponto-Caspian basin but common also in the Baltic Sea was the most common parasite occurring in 65% of N. melanostomus sampled from St Clair River region (Pronin et al., 1997). Recently, N. melanostomus from Lake Erie (Ohio, USA) was found to be a newly described intermediate host for trematode Neochasmus umbellus, the parasite commonly found in piscivorous fishes in the Great Lakes (Kvach and Stepien, 2008) In newly invaded area of the Danube River parasitic load was low and all parasite species found were common. In the Middle Danube basin no parasites parasites specific to Neogobius sp., known from their native population, were observed (Ondrackova et al., 2005).

Means of Movement and Dispersal

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The dispersal of N. melanostomus in Europe may be thought as a combination of natural migration (continuation of postglacial colonization) encouraged by water transport e.g. ships and barges. The human-mediated alteration of river banks e.g. rip-rap habitats and the harbour’s other artificial substrates providing shelter, spawning areas as well as feeding grounds due to presence of zebra mussel beds seem to facilitate N. melanostomus establishment (Ahnelt et al.,1998; Wiesner, 2005). In the case of N. melanostomus invasion of the Baltic Sea ballast-water transport (Sapota, 2004) and possibly as eggs attached to the hulls of barges are the most probable vectors of transmission (Skóra and Stolarski, 1993). The existence of inter-basin connections through artificial canals that join the Black Sea and the Baltic Sea basins enabled N. melanostomus spread. Moreover, as the Danube River system (the Black Sea basin) is connected with rivers of the North Sea basin N. Melanostomus’s further expansion to the rest of Western Europe seems inevitable.

It is thought that the species was transported to the Great Lakes in North America initially in ballast water of ships, which could have come from the Black Sea basin or from the Baltic, and further distributed within the lakes system by similar means (Jude, 1997). Since its first record in 1990, N. melanostomus completed its dispersal throughout the five Great Lakes in 5 years (Jude, 1997).

Pathway Causes

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Cause	Notes	Long Distance	Local	References
Interbasin transfers		Yes		Jude et al. (1992)
Interconnected waterways		Yes	Yes	Pinchuk et al. (2003)

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Ship ballast water and sediment	In all stages of life	Yes	Yes	Corkum et al. (2004); Jude (1997)
Ship hull fouling	Possibly as eggs	Yes	Yes	Corkum et al. (2004); Jude (1997); Jude et al. (1992)

Impact Summary

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Category	Impact
Economic/livelihood	Negative
Environment (generally)	Negative
Human health	Negative

Economic Impact

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In the Baltic Sea N. melanostomus is caught incidentally in eel traps (up to 50 kg/day/boat) but because there is no legalized trade of that species they have no commercial value for fishermen (Skóra, 1996; Charlebois et al., 1997). However, the species is becoming a popular sport fish on the Baltic coast. On the other hand N. melanostomus may negatively interfer with anglers activity as they remove bait from hooks and are caught instead of sport fish. Moreover N. melanostomus is a predator of eggs and juveniles of native game fish species. e.g. in the Great Lakes, reducing the hatching success of lake trout (Salvelinus namaycush) and lake sturgeon (Acipenser fulvescens) (Corkum et al., 2004).

Environmental Impact

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Impact on Habitats

The invasion of N. melanostomus has resulted in the diet shifts among predators and changes in food web structure e.g. in the Gulf of Gdansk (Baltic Sea, Poland), cormorants have shifted their diet from eel (Anguilla anguilla) and sprat (Spratus spratus) to N. melanostomus, resulting in increases in eel and sprat (Bzoma and Stempniewicz, 2001). Increase in planktivorous sprat have lead to a reduction of zooplankton and as a consequence increases in algal biomass have been noted.

As a molluscivore, N. melanostomus has been thought to be of potential use in the Great Lakes basin as a consumer of the introduced zebra mussel (Dreissena polymorpha), which became a serious problem for natural ecosystems and human activities. The zebra mussel is the main dietary component of larger N. melanostomus, while it is not or is rarely consumed by native fish species. The possible reduction of zebra mussel population due to predation is only partly advantageous for ecosystems because the entry of zebra mussel biomass into the food chain of N. melanostomus and their larger fish predators might transfer harmful pollutants into the higher trophic levels (Charlebois et al., 1997).

Kuhns and Berg (1999) reported that predation on invertebrates other than molluscs by N. melanostomus has led to increased algal biomass as measured by chlorophyll content.

Impact on Biodiversity

N. melanostomus negatively affects the recruitment of native fishes in the Great Lakes e.g. it feeds on eggs of lake trout (Salvelinus namaycush) and lake sturgeon (Acipenser fluvescens) probably reducing their hatching success (Corkum et al., 2004). In the St. Clair River the decline of mottled sculpin (Cottus bairdi) coincided with presence of N. melanostomus (Janssen and Jude, 2001). Janssen and Jude (2001) indicated that recruitment failure of mottled sculpins resulted from spawning interference by N. melanostomus. The mechanism of this impact appears to be the aggressive nature of N. melanostomus which drives mottled sculpin from prime feeding, shelter, and especially spawning areas (Janssen and Jude, 2001). Another local species that may be at risk is the deepwater sculpin (Myoxocephalus thompsoni), whose habitats may be penetrated in winter by N. melanostomus (Jude, 1997).

In the southern coasts of the Baltic Sea N. melanostomus may compete for food with flounder (Platichthys flesus) and for space with other native Gobiidae (especially Pomatoschistus microps, Pomatoschistus minutus, Gobius niger) and the eelpout (Zoarces viviparous) (Corkum et al., 2004).

Researchers have suggested a possible link between N. melanostomus. and botulism, Clostridium botulinum type E, a disease of wild migratory birds (Corkum et al., 2004).

Threatened Species

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Threatened Species	Conservation Status	Where Threatened	Mechanism	References	Notes
Villosa fabalis (rayed bean)	EN (IUCN red list: Endangered); National list(s); USA ESA listing as endangered species	USA	Predation	US Fish and Wildlife Service (2012)

Social Impact

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Because N. melanostomus feed on benthic organisms, mainly molluscs, that are exposed to contaminated sediments and are preyed upon by various sport and commercial fishes, the fish diet of humans is a health concern (Corkum et al., 2004).

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
Highly adaptable to different environments
Is a habitat generalist
Pioneering in disturbed areas
Capable of securing and ingesting a wide range of food
Highly mobile locally
Benefits from human association (i.e. it is a human commensal)
Fast growing
Has high reproductive potential
Gregarious
Has high genetic variability

Impact outcomes

Altered trophic level
Conflict
Damaged ecosystem services
Modification of natural benthic communities
Modification of nutrient regime
Negatively impacts human health
Negatively impacts livelihoods
Negatively impacts aquaculture/fisheries
Negatively impacts tourism
Reduced native biodiversity
Threat to/ loss of endangered species
Threat to/ loss of native species

Impact mechanisms

Competition - monopolizing resources
Competition (unspecified)
Interaction with other invasive species
Parasitism (incl. parasitoid)
Predation
Rapid growth

Likelihood of entry/control

Highly likely to be transported internationally accidentally
Highly likely to be transported internationally illegally
Difficult/costly to control

Uses

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Economic Value

N. melanostomus used to be an important component of the commercial catch in the Black Sea and Azov Sea areas. The massive decline in N. melanostomus stocks between the 1960s and 1989 resulted in decline in a commercial fishery (Pinchuk et al., 20003). In newly invaded areas there is no commercial value for N. melanostomus.

Uses List

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Animal feed, fodder, forage

Bait/attractant

Human food and beverage

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

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

Several actions are undertaken to eradicate or at least control and prevent spread of N. melanostomus in the Great Lakes basin. They include: physical activities like using traps, dams and electrical barriers to deter movement (Savino et al., 2001); applying chemicals that are toxic to fish (Dawson et al.,1998); prohibiting transport of N. melanostomus for use as live bait to limit spread; increasing public awareness through publishing educational materials e.g. Sea Grant Program network, Ministry of Natural Resources (Ontario) and other agencies. In 2005, en experimental N. melanostomus eradication project was undertaken in the Pefferlaw Brook, a tributary of Lake Simcoe to eradicate N. melanostomus before it entered Lake Simcoe. Treatment with rotenone was deemed to be the most suitable approach to remove N. melanostomus from the Pefferlaw Brook. The experiment gave positive results as all N. melanostomus were killed (http://www.invadingspecies.com).

References

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Ahnelt H, Banarescu P, Spolwind R, Harka Á, Waidbacher H, 1998. Occurence and distribution of three gobiid species (Pisces, Gobiidae) in the middle and upper Danube region - examples of dispersal patterns? Biologia, Bratislava, 53(5):665-678.

Beek GCWvan, 2006. The round goby Neogobius melanostomus first recorded in the Netherlands. Aquatic Invasions, 1(1):42-43. http://www.aquaticinvasions.ru/2006/AI_2006_1_1_van_Beek.pdf

Bij Vaate Ade, Jazdzewski K, Ketelaars H, Gollasch S, Velde Gder, 2002. Geographical patterns in range expansion of macroinvertebrate Ponto-Caspian species in Europe. Canadian Journal of Fisheries Aquatic Sciences, 59:1159-1174.

Brown JE, Stepien CA, 2008. Ancient divisions, recent expansions: phylogeography and genetics of the round goby Apollonia melanostoma. Mol. Ecol, 17(11):2598-2615.

Butler, R. S., Biggins, R. G., 2004. In: Recovery Plan for Cumberland Elktoe (Alasmidonta atropurpurea), Oyster Mussel (Epioblasma capsaeformis), Cumberlandian Combshell (Epioblasma brevidens), Purple Bean (Villosa perpurpurea), and Rough Rabbitsfoot (Quadrula cylindrica strigillata). US Fish and Wildlife Service, ix + 168 pp..

Bzoma S, Stempniewicz L, 2001. Great cormorants (Phalacrocorax carbo) diet in the Gulf of Gdansk in 1998. In: Third International Symposium on Functioning of Coastal Ecosystems in Various Geographic Regions, 19-22 June 2001, Institute of Oceanography, University of Gdansk.

Charlebois PM, Corkum LD, Jude DJ, Knight C, 2001. The round goby (Neogobius melanostomus) invasion: current research and future needs. Journal of Great Lakes Research, 27(3):263-266.

Charlebois PM, Marsden JE, Goettel RG, Wolf RK, Jude DJ, Rudnika S, 1997. The round goby Neogobius melanostomus (Pallas, 1811), a review of European and North American Literature. Illinois Natural History Survey Special Publication., 1-76.

Corkum LD, Sapota MR, Skora KE, 2004. The round goby, Neogobius melanostomus, a fish invader on both sides of the Atlantic Ocean. Biological Invasions, 6(2):173-181.

Dawson VK, Boogaard MA, Bills TD, 1998. Evaluation of piscicides for controlling range expansion of round goby (Neogobius melanostomus) and ruffe (Gymnocephalus cernuus). United States Geological Survey, Biological Resource Division.

Dillon AK, Stepien CA, 2001. Genetic and biogeographical relationships of the invasive round (Neogobius melanostomus) and tubnose (Proterorhinus marmoratus) gobies in the Great Lakes versus European populations. Journal of the Great Lakes Research, 27:267-280.

Dobrovolov JS, Georgiiev TU, Dobrovolova SG, 1995. Comparative electrophoretic investigation of the species of the family Gobiidae (Pisces) in the Bulgarian sector of the Black Sea. Proceedings of the Institute of Fisheries (Varna), 23:48-68.

Dougherty JD, Moor WS, Ram JL, 1996. Mitochondrial DNA analysis of round goby (Neogobius melanostomus) and tubnose goby (Proterorhinus marmoratus) in the Great Lakes basin. Canadian Journal of Fisheries and Aquatic Sciences, 53:474-480.

Gulugin SY, Kunitsky DF, 1999. New data on spread genus Neogobius: N. fluviatilis, N. melanostomus, N. gymnotrachelus. In: Thesis of International Scientific Conference, Kaliningrad,.

Guti G, Eros T, Szalóky Z, Tóth B, 2003. Round goby, Neogobius melanostomus (pallas, 1811) in the Hungarian section of the Danube. Halászat, 93(3):116-119.

Janssen J, Jude DJ, 2001. Recruitment failure of mottled sculpin Cottus bairdi in Calumet Harbor, southern Lake Michigan, induced by the newly introduced round goby Neogobius melanostomus. Journal of Great Lakes Research, 27(3):319-328.

Jude DJ, 1997. Round goby: cyberfish of the third millennium. Great Lakes Research Review, 3(1):27-34.

Jude DJ, Reider RH, Smith GR, 1992. Establishment of Gobiidae in the Great Lakes basin. Canadian Journal of Fisheries and Aquatic Sciences, 49:416-421.

Jurajda P, Cerny J, Polacik M, Valová Z, Janác M, Blažek R, Ondracková M, 2005. The recent distribution and abundance of non-native Neogobius fishes in the Slovak section of the river Danube. Journal of Applied Ichthyology, 21(4):319-323.

Kostrzewa J, Grabowski M, 2002. Monkey goby, Neogobius fluviatilis (Pallas, 1811), In the Vistula River - phenomenon of Ponto-Caspian Gobiidae. Przeglad Zoologiczny, 46(3-4):235-242.

Kuhns LA, Berg MB, 1999. Benthic invertebrate community responses to round goby (Neogobius melanostomus) and zebra mussel (Dreissena polymorpha) iinvasion in southern Lake Michigan. Journal Great Lakes Research, 25:910-971.

Kuhns LA, Nekrasova MY, Revina NI, 1974. On the diet of round goby (Neogobius melanostomus) and utilization of food supply in the Azov Sea. Russian Journal of Zoology, 53:728-736.

Kvach Y, Skóra KE, 2007. Metazoa parasites of the invasive round goby Apollonia melanostoma (Neogobius melanostomus) (Pallas) (Gobiidae: Osteichthyes) in the Gulf of Gdan´sk, Baltic Sea, Poland: a comparison with the Black Sea. Parasitology Research, 100(4):767-774. http://www.springerlink.com/link.asp?id=100447

Kvach Y, Stepien CA, 2008. The invasive round goby Apollonia melanostoma (Actinopterygii: Gobiidae) - a new intermediate host of the trematode Neochasmus umbellus (Trematoda: Cryptogonimidae) in Lake Erie, Ohio, USA. Journal of Applied Ichthyology, 24(1):103-105. http://www.blackwell-synergy.com/loi/jai

Moskalkova KI, 1996. Ecological and morphophysiological prerequisites to range extention in the round goby Neogobius melanostomus under conditions of anthropogenic pollution. Journal of Ichthyology, 36:584-590.

Nelson JS, 1994. Fishes of the world. Fishes of the world., Ed. 3:xvii + 600 pp.

Ojaveer H, 2006. The round goby Neogobius malanostomus is colonizing the NE Baltic sea. Aquatic Invasions, 1:44-45.

Ondracková M, Dávidová M, Pecínková M, Blazek R, Gelnar M, Valová Z, Cerný J, Jurajda P, 2005. Metazoan parasites of Neogobius fishes in the Slovak section of the River Danube. Journal of Applied Ichthyology, 21(4):345-349. http://www.blackwell-synergy.com/servlet/useragent?func=showIssues&code=jai

Pinchuk VI, Vasil'eva ED, Vasil'ev VP, Miller P, 2003. Neogobius melanostomus (Pallas, 1811). In: The Freshwater Fishes of Europe. Vol. 8/I Mugilidae, Atherinidae, Atherionopsidae, Blennidae, Odontobutidae, Gobiidae 1 [ed. by Miller PJ]: AULA-Verlag, 293-345.

Pronin NM, Fleischer GW, Baldanova DR, Pronina SV, 1997. Parasites of the recently established round goby (Neogobius melanostomus) and tubenose goby (Proterorhinus marmoratus) (Cottidae) from the St. Clair River and Lake St. Clair, Michigan, USA. Folia Parasitologica, 44(1):1-6.

Rolbiecki l, 2006. Parasites of the round goby, Neogobius melanostomus (Pallas, 1811), an invasive species in the Polish fauna of the Vistula Lagoon ecosystem. Oceanologia, 48(4):545-561.

Sapota MR, 2004. The round goby (Neogobius melanostomus) in the Gulf of Gdansk - a species introduction into the Baltic Sea. Hydrobiologia, 514:219-224.

Sapota MR, Skora KE, 2005. Spread of alien (non-indigenous) fish species Neogobius melanostomus in the Gulf of Gdansk (south Baltic). Biological Invasions, 7:157-164.

Savino JF, Jude DJ, Kostich MJ, 2001. Use of electrical barriers to deter movement of round goby. In: American Fisheries Society Symposium, Bethesda, Maryland, 171-182.

Simonovic P, Valkovic B, Paunovic M, 1998. Round goby Neogobius melanostomus, a new Ponto-Caspian element for Yugoslavia. Folia Zoological, 47:305-312.

Skóra KE, 1996. Report of the working group on introduction and transfers of marine organisms (WGITMO). In: ICES Annual Science Conference, Reykjavik, Iceland, 96-107.

Skóra KE, Stolarski J, 1993. New fish species in the Gulf of Gdansk. Neogobius sp [cf Neogobius melanostomus (Pallas 1811)]. Bulletin of the Sea Fisheries Institute, Gdynia, 1(128):83.

Smirnov AI, 1986. Perciformes (Gobioidei), Scorpaeniformes, Pleuronectiformes, Gobiesociformes, Lophiiformes. Fauna of Ukraine 8. Fishes (5). Naukova Dumka, Kiev.

Stránai I, Andreji J, 2004. The first report of round goby, Neogobius melanostomus (Pisces, Gobiidae) in the waters of Slovakia. Folia Zoologica, 53(3):335-338.

US Fish and Wildlife Service, 2012. In: Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for the Rayed Bean and Snuffbox Mussels Throughout Their Ranges; Final Rule. 77(30) US Fish and Wildlife Service, 8632-8665. https://www.gpo.gov/fdsys/pkg/FR-2012-02-14/pdf/2012-2940.pdf

US Fish and Wildlife Service, 2013. In: Endangered and Threatened Wildlife and Plants; Determination of Endangered Species Status for 15 Species on Hawaii Island; Final Rule. 78(209) US Fish and Wildlife Service, 64638-64690. https://www.gpo.gov/fdsys/pkg/FR-2013-10-29/pdf/2013-24103.pdf

Wiesner C, 2005. New records of non-indigenous gobies (Neogobius spp.) in the Austrian Danube. Journal of Applied Ichthyology, 21:324-327.

Wiesner C, Spolwind R, Waidbacher H, Guttmann S, Doblinger A, 2000. [English title not available]. (Erstnachweis der Schwarzmundgrundel Neogobius melanostomus (Pallas, 1814) in Österreich) Österreichs Fischerei, 53:330-331.

Distribution References

Beek G C W van, 2006. The round goby Neogobius melanostomus first recorded in the Netherlands. Aquatic Invasions. 1 (1), 42-43. DOI:10.3391/ai.2006.1.1.10

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

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

Corkum L D, Sapota M R, Skora K E, 2004. The round goby, Neogobius melanostomus, a fish invader on both sides of the Atlantic Ocean. Biological Invasions. 6 (2), 173-181.

Gulugin S Y, Kunitsky D F, 1999. New data on spread genus Neogobius: N. fluviatilis, N. melanostomus, N. gymnotrachelus. In: International Scientific Conference, Kaliningrad, Vol. 1, Pt. 5 [International Scientific Conference, Kaliningrad, Vol. 1, Pt. 5],

Guti G, Eros T, Szalóky Z, Tóth B, 2003. Round goby, Neogobius melanostomus (pallas, 1811) in the Hungarian section of the Danube. ([Hungarian title not available]). Halászat. 93 (3), 116-119.

Jude D J, 1997. Round goby: cyberfish of the third millennium. Great Lakes Research Review. 3 (1), 27-34.

Jude D J, Reider R H, Smith G R, 1992. Establishment of Gobiidae in the Great Lakes basin. Canadian Journal of Fisheries and Aquatic Sciences. 416-421.

Ojaveer H, 2006. The round goby Neogobius malanostomus is colonizing the NE Baltic sea. Aquatic Invasions. 44-45.

Pinchuk V I, Vasil'eva E D, Vasil'ev V P, Miller P, 2003. Neogobius melanostomus (Pallas, 1811). In: The Freshwater Fishes of Europe. Vol. 8/I Mugilidae, Atherinidae, Atherionopsidae, Blennidae, Odontobutidae, Gobiidae 1. [ed. by Miller P J]. AULA-Verlag. 293-345.

Seebens H, Blackburn T M, Dyer E E, Genovesi P, Hulme P E, Jeschke J M, Pagad S, Pyšek P, Winter M, Arianoutsou M, Bacher S, Blasius B, Brundu G, Capinha C, Celesti-Grapow L, Dawson W, Dullinger S, Fuentes N, Jäger H, Kartesz J, Kenis M, Kreft H, Kühn I, Lenzner B, Liebhold A, Mosena A (et al), 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. 8 (2), 14435. http://www.nature.com/articles/ncomms14435

Simonovic P, Valkovic B, Paunovic M, 1998. Round goby Neogobius melanostomus, a new Ponto-Caspian element for Yugoslavia. Folia Zoologica. 305-312.

Skóra K E, Stolarski J, 1993. New fish species in the Gulf of Gdansk. Neogobius sp. [cf. Neogobius melanostomus (Pallas 1811)]. Bulletin of the Sea Fisheries Institute, Gdynia. 1 (128), 83.

Wiesner C, Spolwind R, Waidbacher H, Guttmann S, Doblinger A, 2000. First record of Neogobius melanostomus (Pallas, 1814) in Austria. (Erstnachweis der Schwarzmundgrundel Neogobius melanostomus (Pallas, 1814) in Österreich). Österreichs Fischerei. 330-331.

Links to Websites

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Website	URL	Comment
CRS Report for Congress: Harmful Non-Native Species	http://www.nsceonline.org/
Culprit-Great Lakes Goby	http://www.culprit.com/html/great_lakes_goby.html
Fish of Great Lakes by the Wisconsin Sea Grant	http://www.seagrant.wisc.edu/greatlakesfish
The North European and Baltic Network on Invasive Alien Species	http:// www.nobanis.org/

Contributors

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28/04/08 Original text by:

Joanna Grabowska, University of Lodz, Dept Invertebrate Zoology & Hydrobiology, Banacha 12/16, 90-237 Lodz, Poland

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Neogobius melanostomus (round goby)

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