NOBANIS – Invasive Alien Species Fact Sheet

Dikerogammarus villosus

Authors of this fact sheet:  Vitaliy Semenchenko, Laboratory Hydrobiology, Scientific and Practical Centre for Bioresources, National Academy of Sciences, Republic of Belarus, e-mail: [email protected]  Simon Devin, LIEC, Université de Lorraine, UMR CNRS 7360, France, e-mail: [email protected]

Bibliographical reference – how to cite this fact sheet: Semenchenko, V. Devin, S (2017): NOBANIS – Invasive Alien Species Fact Sheet – villosus. – From: Online Database of the European Network on Invasive Alien Species – NOBANIS www.nobanis.org, Date of access x/x/201x.

Species description

Scientific names: (Sowinsky, 1894)

Synonyms: none

Common names: killer shrimp (English), Großer Höckerflohkrebs (German)

Taxonomy Phylum - Arthropoda Class - Crustacea Order - Family -

Identification The body size of adult specimens is up to 30 mm. D. villosus can be recognized by clearly marked tubercules on urosome segments I and II with 3-5 spines. (Mordukhai-Boltowskoi 1960; Dobson, 2002, see Photo). The closely related species Dikerogammarus bispinosus Martynov, 1925 has only two spines. The second antennae has a sparsely haired peduncle and a flagellum with dense setae (MacNeil et al, 2010). Large and powerful mandibles characterize D. villosus as an effective predator (Mayer et al, 2008)

Photo: A. Makarenko

Similar Species: Dikerogammarus bispinosus Martynov, 1925

For further information:  GLANSIS: https://nas.er.usgs.gov/queries/greatlakes/FactSheet.aspx  DAISIE: http://www.europe-aliens.org/pdf/Dikerogammarus_villosus.pdf  CABI: http://www.cabi.org/isc/datasheet/108309

Native distribution Ponto-Caspian basin. The species is typical in estuaries of the Dnieper, Don, Dnister, Volga and rivers and in the limans and lagoons in Ukraine, Romania and Bulgaria (Mordukhai-Boltowskoi 1960; Dedyu, 1980).

History of introduction and geographical spread D. villosus is native in the estuaries of the Volga, Don, Dnister, Danube and Dnieper rivers. But in the middle and upper reaches of these rivers it is alien. D. villosus penetrated European water bodies through man-made canals. The migration routes of D. villosus are the central corridor connecting the Dnieper – Prypiat - Vistula rivers, and the southern corridor connecting the Danube and rivers (Ludwig canal), and canals connecting Oder and Elbe rivers (Bij de Vaate et al, 2002; Grabowski et al, 2007; Rewicz et al, 2014). In southern corridor D. villosus is native in Lower Danube River, but alien for the Upper and Middle Danube River. According to Bij de Vaate et al (2002) and Rewicz et al (2014) the species penetrated the Main and Oder rivers and afterwards France inland waters via southern corridor (Main–Danube Canal). Introduction of D.villosus to some Alpine lakes may be associated with the recreational boating and scuba-diving activity (Bacela-Spychalska et al, 2013). Since the 1960’s D.villosus has been successfully introduced to a number of Ukrainian artificial dam reservoirs (Semenchenko et al, 2016). After that, the killer shrimp spread via central corridor to the Vistula River.

Distribution in the European countries D. villosus was first found in the central corridor (Prypiat River, Vistula River) in 2007 (Bacela et al, 2008; Semenchenko et al, 2009), in the upper Danube in 1992 by Tittizer et al (2000), and it penetrated into the Dutch part of the Lower Rhine River via the Main–Danube Canal in 1994 (Bij de Vaate and Klink, 1995). It was recorded for the first time in the Czech Republic in 2003 (Petrusek, Beran, 2006) and in the French and Slovakia inland waters in 1997 (Devin et al, 2001; Šporka, 1999), Switzerland in about 1990 (Altermatt et al, 2014), Hungary, including Lake Balaton (Musko, 1989), Croatia in 2007 (Žganec, Gottstein, Hudina, 2009), Lithuania in 2015 (Šidagytė et al, 2017) and British waters in 2010 (MacNeil et al, 2010). In the middle Volga River this species first found in 2000s (Yakovlev, Yakovleva, 2005). Nowadays killer shrimp occurs in 20 European countries (see Table I).

Country Not found Local Common Austria x Belarus x Belgium x Bulgaria* x Croatia x Czech Republic x Denmark x Estonia x European part of x Russia* Finland x France x x Greenland x Ireland x Italy x Hungary x Latvia x Lithuania x Netherlands x Norway x Poland x Romania* x Serbia and x Montenegro Sweden x Switzerland x Slovakia x United Kingdom x Ukraine* x Table 1. Distribution D.villosus in Europe *- native in the Black Sea estuarias, limans and lagoons Local – recorded in several localities Common – recorded in many localities

Alien status in region This species is invasive in the European countries.

Pathways and vectors Shipping (fouling and ballast waters) and natural spread upstream the Danube, Rhine, Dnieper, Don rivers and other large rivers of Central and Western Europe via canals connecting various river drainages (invasive corridors) (Panov et al, 2009; Gherardi et al, 2009; Rewicz et al, 2014). This species has a high ability to remain attached to objects, such as boat, and recreational boating equipment. It has ability to survive harsh transport conditions and being out of water for up to 3-6 (in shell clusters) days (Bacela-Spychalska et al, 2013; Rewicz et al, 2014). The spread D.villosus in Danube River occured by active movement and drift or as a result of ship transport (Paunovic et al, 2015). The spread of D.villosus along the inshore Baltic waters likely occurred with fouling organisms, as well as by natural migration (Jazdzewski et al, 2002). The pathways of introduction to the British waters are unknown (MacNeil et al, 2010), but DNA analysis showed that the UK populations are similar to populatitons from southen corridor and genetically distinct from the central route (Rewicz et al, 2015). Killer shrimp was not recorded in

Ecology This species has wide environmental tolerances, to tolerate temperatures from 0-35°C, with an optimal temperature range of 20-23°C (Bruijs et al, 2001; van der Velde et al, 2009; Wijnhoven et al, 2003). D. villosus can live at salinity up to 12‰ and is able to adapt to salinities of up to 20‰. No correlations between the abundance of species and water hydrochemistry were found (Bacela-Spychalska et al, 2013; Semenchenko et al, 2013). Killer shrimp can tolerate even lower oxygen concentrations (letal concentration is 0.35 mg O2/l, Dedyu, 1980). It prefers coastal waters with vegetated areas with Potamogeton sp. and Ceratophyllum sp., mats of algae near or on the water surface, occurred on stony substratum and zebra mussel beds, occasionally on sand (Devin and Beisel, 2006; Devin et al, 2003; Semenchenko et al, 2013). This species reaches the highest densities on hard substrates, primarily boulders, rocks, and pebbles (Kley, Maier, 2005; Platvoet et al, 2009).

Reproduction This amphipod reproduces all year round in its native range (Mordukhai-Boltovskoi, 1949). In the new area this species is characterized by a long reproductive period, early sexual maturity, short generation and life span time, short duration of embryonic development (Pockl, 2009). The mature females are approximately 6 mm (Devin et al, 2004). They start reproducing when the water temperature increases above 13°C, and ovigerous females mainly occur from April to October (Devin et al, 2004). However, some individuals are still sexually active when water temperature is colder (Pockl, 2007). Females of D. villosus produce large clutches; mean clutch size during the reproduction period ranges from 29 to 195 eggs per ovigerous female (Kley, Maier, 2003; Pockl, 2007).

Impact D. villosus is one of a hundred of the worst invasive alien species in Europe (DAISIE, 2009). The killer shrimp is an omnivorous species. It kills its preys by biting and shredding them. Interactions (competition and predation; Dick, Platvoet, Kelly, 2002; Kinzler et al, 2009) between D. villosus and native gammarid species can result in displacement or local of native species, thereby reducing biodiversity (Kley, Maier, 2005; Semenchenko et al, 2013). The predation of D.villosus leads to altertion in macroinvertebrate assemblages. D. villosus has also been observed attacking small fish and its spawn (Casellato, Visentin, La Piana, 2007) and thereby have exerted the impact on food webs (van Riel et al, 2006). Besides, this species is a ”reservoir” for different pathogens such as Microsporidia, Trematoda and others (Bojko et al, 2013).

Management To avoid the spread of the killer shrimp through boating activities, recommended management measures include inspecting and cleaning boats both before and after use, and draining bilge water from boats before leaving any given site (Anderson et al, 2014). Besides, fishing waders, fish nets, other equipment of anglers and canoeists must be checked (Check, Clean, Dry system) (GB non-native species secretariat, 2017). The ballast water can be treated (DAISIE, 2009).

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