Dikerogammarus Villosus
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''Dikerogammarus villosus'', also known as the killer shrimp, is a species of amphipod
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
native to the Ponto-Caspian region of eastern Europe, but which has become invasive across the western part of the continent. In the areas it has invaded, it lives in a wide range of habitats and will prey on many other animals. It is fast-growing, reaching sexual maturity in 4–8 weeks. As it has moved through Europe, it threatens other species and has already displaced both native amphipods and previous invaders.


Description

''D. villosus'' can grow up to in length, relatively large for a freshwater amphipod. It varies in appearance, with some specimens being striped, and some not. It has relatively large
mandibles In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
which allow it to be an effective predator.


Distribution

''D. villosus'' was originally found in the lower courses of large rivers in the
Black Sea The Black Sea is a marginal mediterranean sea of the Atlantic Ocean lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bounded by Bulgaria, Georgia, Rom ...
and
Caspian Sea The Caspian Sea is the world's largest inland body of water, often described as the world's largest lake or a full-fledged sea. An endorheic basin, it lies between Europe and Asia; east of the Caucasus, west of the broad steppe of Central Asia ...
drainage basin A drainage basin is an area of land where all flowing surface water converges to a single point, such as a river mouth, or flows into another body of water, such as a lake or ocean. A basin is separated from adjacent basins by a perimeter, ...
s. It has become an invasive species across central and western Europe, using the
Danube The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , p ...
river and its
tributaries A tributary, or affluent, is a stream or river that flows into a larger stream or main stem (or parent) river or a lake. A tributary does not flow directly into a sea or ocean. Tributaries and the main stem river drain the surrounding drainag ...
in its expansion. It is thought to have first escaped from the Danube in 1992 when the
Rhine–Main–Danube Canal The Rhine–Main–Danube Canal (German: ''Rhein-Main-Donau-Kanal''; also called Main-Danube Canal, RMD Canal or Europa Canal), is a canal in Bavaria, Germany. Connecting the Main and the Danube riv ...
was opened and has since spread to nearly all the major rivers in western Europe, including the
Rhône The Rhône ( , ; wae, Rotten ; frp, Rôno ; oc, Ròse ) is a major river in France and Switzerland, rising in the Alps and flowing west and south through Lake Geneva and southeastern France before discharging into the Mediterranean Sea. At Ar ...
,
Loire The Loire (, also ; ; oc, Léger, ; la, Liger) is the longest river in France and the 171st longest in the world. With a length of , it drains , more than a fifth of France's land, while its average discharge is only half that of the Rhône ...
, Seine, Moselle,
Meuse The Meuse ( , , , ; wa, Moûze ) or Maas ( , ; li, Maos or ) is a major European river, rising in France and flowing through Belgium and the Netherlands before draining into the North Sea from the Rhine–Meuse–Scheldt delta. It has a t ...
,
Rhine ), Surselva, Graubünden, Switzerland , source1_coordinates= , source1_elevation = , source2 = Rein Posteriur/Hinterrhein , source2_location = Paradies Glacier, Graubünden, Switzerland , source2_coordinates= , so ...
and
Main Main may refer to: Geography * Main River (disambiguation) **Most commonly the Main (river) in Germany * Main, Iran, a village in Fars Province *"Spanish Main", the Caribbean coasts of mainland Spanish territories in the 16th and 17th centuries ...
, as well as the
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain. The sea stretches from 53°N to 66°N latitude and ...
. It is unknown how the species is dispersed but it is probably related to shipping activity. It spread quickly through western Europe; found in the Rhine at the German–Dutch border (1995), the canals and rivers of northern Germany (1998), the Baltic
Szczecin Lagoon Szczecin Lagoon ( pl, Zalew Szczeciński, german: Stettiner Haff, since 1945 sometimes also ''Oderhaff'' (Oder lagoon) or ''Pommersches Haff'' (Pomeranian lagoon)) is a lagoon in the Oder estuary, shared by Germany and Poland. It is separated f ...
(2001), the Moselle (2001), the Netherlands (2002), Lake Constance (2003),
Lake Leman , image = Lake Geneva by Sentinel-2.jpg , caption = Satellite image , image_bathymetry = , caption_bathymetry = , location = Switzerland, France , coords = , lake_type = Glacial la ...
, the Rhine in France (2003), the (2003) and
Lake Garda Lake Garda ( it, Lago di Garda or ; lmo, label= Eastern Lombard, Lach de Garda; vec, Ƚago de Garda; la, Benacus; grc, Βήνακος) is the largest lake in Italy. It is a popular holiday location in northern Italy, about halfway between ...
(2003). Its spread is thought to be related to the previous introduction of the
zebra mussel The zebra mussel (''Dreissena polymorpha'') is a small freshwater mussel. The species originates from the lakes of southern Russia and Ukraine, but has been accidentally introduced to numerous other areas and has become an invasive species in ma ...
, ''Dreissena polymorpha'' which it has evolved alongside. In September 2010, it was found in
Grafham Water Grafham Water is an biological Site of Special Scientific Interest (SSSI) north of Perry, Huntingdonshire. It was designated an SSSI in 1986. It is a reservoir with a circumference of about , is deep at maximum, and is the eighth largest rese ...
in
Cambridgeshire Cambridgeshire (abbreviated Cambs.) is a county in the East of England, bordering Lincolnshire to the north, Norfolk to the north-east, Suffolk to the east, Essex and Hertfordshire to the south, and Bedfordshire and Northamptonshire to the ...
, the first report of the species in the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the European mainland, continental mainland. It comprises England, Scotlan ...
and it was found in Wales in November 2010. There are fears that it could spread to the
Great Lakes The Great Lakes, also called the Great Lakes of North America, are a series of large interconnected freshwater lakes in the mid-east region of North America that connect to the Atlantic Ocean via the Saint Lawrence River. There are five lak ...
in North America in future, carried in the
ballast water Ballast is used in ships to provide moment to resist the lateral forces on the hull. Insufficiently ballasted boats tend to tip or heel excessively in high winds. Too much heel may result in the vessel capsizing. If a sailing vessel needs to voy ...
of ships.


Ecology

In its natural range, ''D. villosus'' is not the most abundant species of amphipod and it does not behave as aggressively as it does in areas it has invaded.


Habitat

''D. villosus'' can colonise many types of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
as it is able to tolerate a wide range of temperatures (), low oxygen concentrations, and salinity up to 20 %. It is found in lakes, canals and rivers living in a range of substrates. When found in lotic environments, the species prefers habitats with relatively mild currents. It is thought that zebra mussels change habitats by increasing the amount of benthic organic matter, which benefits ''D. villosus'' helping them to outcompete other species. When given a choice, ''D. Villosus'' spend more time feeding around zebra mussel shells than a bare substrate.


Feeding

''D. villosus'' is
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
and feeds on a variety of
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s, including other members of the
Gammaridae Gammaridae is a family of amphipods. In North America they are included among the folk taxonomic category of " scuds", and otherwise gammarids is usually used as a common name. They have a wide distribution, centered on Eurasia, and are eury ...
family. It has been found to kill
blue-tailed damselfly The blue-tailed damselfly or common bluetail (''Ischnura elegans'') is a damselfly, belonging to the family Coenagrionidae. Subspecies and varieties Subspecies and varieties include: *''Ischnura elegans ebneri'' Schmidt, 1938 *''Ischnura eleg ...
nymphs, water hoglice, water boatman, fish leeches as well as small fish and the eggs of other
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
s. ''D. villosus'' sometimes kills prey but does not eat it; it kills its prey by biting it with its large mandibles and then shreds it before eating it. ''D. villosus'' also consumes many other forms of food, including grazing on biofilms, consuming plant detritus, eating and re-digesting feces, and feeding on organic matter and algae suspended in the water column.


Growth and reproduction

''D. villosus'' breeds all year round so long as the water temperature is above . When they mate, the female is carried on the ventral side of the male. Each pair produces an average of 27 eggs, but up to 50 eggs can be laid. The young animals become sexually mature in 4 to 8 weeks, once they are 6 mm in length and after moulting several times. They are fast-growing, during winter increasing by 1.3–2.9 mm in length per month and by 2.0–2.6 mm every two weeks in spring. Populations are predominantly female.


Effects on other species

In the Netherlands, ''D. villosus'' is threatening the native amphipod species ''
Gammarus duebeni ''Gammarus'' is an Amphipoda, amphipod crustacean genus in the family (biology), family Gammaridae. It contains more than 200 described species, making it one of the most species-rich genera of crustaceans. Different species have different optimu ...
'', as well as '' Gammarus tigrinus'' which had previously become invasive after previously being introduced from North America. It is thought to have displaced two other species of ''
Dikerogammarus ''Dikerogammarus'' is a genus of amphipod crustaceans, containing the following species: *'' Dikerogammarus aralychensis'' (Birstein, 1932) *'' Dikerogammarus batalonicus'' Ponyi, 1955 *'' Dikerogammarus bispinosus'' Martynov, 1925 *'' Dikeroga ...
'' (''D. bispinosus'' and ''D. haemobaphes'') which were previously invasive in the Danube. Its ability to attack and feed on a range of species could cause the local
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
of some species.


References


External links


Species Profile - Killer Shrimp (''Dikerogammarus villosus'')
National Invasive Species Information Center,
United States National Agricultural Library The United States National Agricultural Library (NAL) is one of the world's largest agricultural research libraries, and serves as a national library of the United States and as the library of the United States Department of Agriculture. Located ...
.
Invasive Species Compendium
CAB International {{Taxonbar, from=Q447913 Gammaridae Crustaceans described in 1894