Checklist of Cryptogenic and Alien Crustacea of the European Atlantic Coast

Abstract

An overview is presented of 49 species of alien and cryptogenic marine and estuarine crustaceans established along the Atlantic coast of Europe. The alien species include Diplostraca (1 species), Ostracoda (1), Copepoda (10), Cirripedia (10), Mysidacea (1), Amphipoda (12), Tanaidacea (1), Isopoda (4), and Decapoda (9). The established introductions are primarily from Indo-Pacific, Atlantic America and Mediterranean-Ponto-Caspian regions, and the primary vectors that have brought these species to Europe are ballast water, ship fouling and mariculture activity.

Appendices

Appendix AInventory of Alien and Cryptogenic Species on the European Atlantic Coast

Unless otherwise stated, ecological, environmental, economic or other impacts are unknown; the absence of such reports should not be taken as the absence of impacts.

Branchiopoda Diplostraca (“Cladocera”)

Sidiidae

Penilia avirostris Dana, 1849

A marine cladoceran. Native to Eurasia? Sub-cosmopolitan: Mediterranean Sea and Black Sea (Russia, Bulgaria, Romania, Turkey, Ukraine, Georgia), Red Sea, China. Introduced to Canada, and possibly elsewhere in the cited range. North-East Atlantic: introduced to The Netherlands and North Sea (Carlton 1985; Johns et al. 2005) but said to be native to Germany (Greve et al. 2004). Biology/ecology: marine, brackish, fresh. Vector(s): possibly ballast water. Status: Cryptogenic.

Ostracoda

Sarsiellidae

Eusarsiella zostericola (Cushman, 1906) [Sarsiella zostericola]

A marine ostracod. Native to the USA Atlantic coast. North-East Atlantic: Introduced to South-East English estuaries by the end of nineteenth or first half of twentieth century (Bamber 1987a, b; Eno et al. 1997). Biology/ecology: see Bamber 1987a, b. Vector(s): Importations of the American oyster Crassostrea virginica (Eno et al. 1997). Comments: Information on the occurrence of this very small species in other European localities is limited. Status: Introduced.

Copepoda

Acartiidae

Acartia omorii Bradford, 1976 [Acartia (Acartiura) omorii]

A calanoid copepod. Native to Japan-China. Introduced to California and Chile. North-East Atlantic: Dunkerque, France (Brylinski 2009; Razouls et al. 2009), North Sea (Seuront 2005). Biology/ecology: epipelagic. Vector(s): ballast water (Razouls et al. 2009). Status: Introduced.

Acartia tonsa Dana, 1849 [Acartia tonsa cryophylla]

An estuarine copepod. Cosmopolitan, native distribution unknown, possibly Indo-Pacific. North-East Atlantic range: First reported from France (Rémy 1927), but first European records are from Dutch waters about 1912–1916 (Redeke 1935); also occurring in UK, Belgium and Denmark (Polk 1963; Brylinski 1981; Eno et al. 1997; Reise et al. 1999; Wolff 2005). Biology/ecology: Tolerant to low salinities. Diapause eggs present (Zilhoux and Gonzalez 1972). Vector(s): ballast water. Status: Cryptogenic.

Temoridae

Eurytemora americana Williams, 1906

An estuarine copepod. Native to North-East Pacific (Canada, Alaska, Oregon, USA). Introduced to Argentina (Hoffmeyer et al. 2000). North-East Atlantic: Iceland: (Jespersen 1940), Oosterschelde, The Netherlands (Bakker 1994; Wolff 2005), English Channel, Port of Dunkerque, Straight of Dover (Razouls et al. 2009). Biology/ecology: Brackish ponds; coastal waters. Vector(s): Ballast water. Status: Introduced.

Eurytemora pacifica Sato, 1913

A calanoid copepod. Native to North Pacific (China, Korea, Japan, Alaska). North-East Atlantic range: France only Charente estuary (Goulletquer et al. 2002, 2004). Biology/ecology: marine and brackish waters. Vector(s): Ballast water. Status: Introduced.

Myicolidae

Midicola spinosus (Raffaele and Monticelli, 1885) [Pseudomyicola spinosus]

A parasitic copepod. Native to Pacific, widely distributed, Japan to Mexico. Introduced (possibly) to Indian Ocean: Madagascar and the Atlantic: North Carolina, Bermuda, West Indies, Brazil, Mediterranean Sea, Black Sea (Humes 1968). North-East Atlantic: Arcachon Bay, France (Goulletquer et al. 2002). Biology/ecology: optional parasitic. Vector(s): Mariculture. Status: Introduced.

Myicola ostreae Hoshina and Sugiura, 1953

A parasitic copepod. Native to North-West Pacific (Japan, Korea). North-East Atlantic: The Netherlands, Ireland, France (Comps 1972; Reise et al. 1999; Goulletquer et al. 2002, 2004; Wolff 2005). Biology/ecology: parasitic in oysters. Vector(s): Mariculture: Accidentally introduced into France with imports of Crassostrea gigas from Japan in the 1970s (His 1979), since then has been found in several European countries (Streftaris et al. 2005). Status: Introduced.

Myticolidae

Mytilicola intestinalis Steuer, 1902

A parasitic copepod. Native to Mediterranean. North-East Atlantic: Established in Denmark, The Netherlands, Germany, Belgium (Leloup and Lefevere 1952; Polk 1963), Ireland, UK, France (Davey 1989; Goulletquer et al. 2002, 2004; Wolff 2005; Kerckhof et al. 2007). Biology/ecology: Parasitic in mussels. Impacts: Relatively harmless (Dare 1985). Vector(s): Mariculture. Status: Introduced.

Mytilicola orientalis Mori, 1935

A parasitic copepod. Native to North-West Pacific. Introduced to the Pacific coast of Canada and the United States. North-East Atlantic: France (His 1977, 1979), The Netherlands, Ireland, UK (Wolff 2005). Biology/ecology: Parasitic in Crassostrea gigas, Mytilus edulis, Ostrea edulis (Stock 1993a, b). Vector(s): Mariculture. Accidentally introduced into Europe with Pacific oysters (Stock 1993a). Status: Introduced.

Porcellidiidae

Porcellidium ovatum Haller, 1879

A harpacticoid copepod. Native region unclear: first described from Italy, and since recorded from Kenya, Indonesia, and the Caribbean Sea. North-East Atlantic: Ireland, UK and surrounding seas (Holmes et al. 1997; Minchin and Eno 2002; Stokes et al. 2004). Biology/ecology: estuaries. Vector(s): possibly ballast water or ballast sediments or fouling on ships’ hulls. Status: Cryptogenic.

Spiophanicolidae

Spiophanicola spinosus Ho, 1984 [Spiophanicola spinulosus Ho, 1984]

A poecilostomatid copepod. Possibly native to California, but perhaps circumboreal. North-East Atlantic: introduced to Cullercoats, Scotland (O’Reilly 1999). Biology/ecology: associated with the polychaete Spiophanes kroyeri. Vector(s): possibly ballast water or ballast sediments. Status: Cryptogenic.

Cirripedia

Archeobalanidae

Austrominius modestus (Darwin, 1854) [Elminius modestus]

An Australasian barnacle. Native to Australia, New Zealand. North-East Atlantic: First specimens found in Portsmouth, England in 1943 (Stubbings 1950), then extended range progressively. This species is now found in Ireland, Shetland Islands, Denmark, The Netherlands, Germany to Gibraltar and Madeira (Crisp 1958; Eno et al. 1997; Wolff 2005; Minchin 2007). Biology/ecology: Rapid growth and tolerant to turbid waters with low salinity. Reaches maturity by first year, several broods per year. Impacts: Competes with other barnacles such as Balanus spp. and Chthamalus spp. (Eno et al. 1997; Kerckhof 2002; Kerckhof et al. 2007). Vector(s): pelagic larvae in ballast waters, fouling on ships’ hulls. Comments: Established along most European Atlantic coastlines where it is fairly common on several intertidal hard substrates. Status: Introduced.

Balanidae

Amphibalanus amphitrite Darwin, 1854

The striped or purple acorn barnacle. Native to Tropical Seas: South-western Pacific and Indian Oceans (according to fossil records), and introduced to North Pacific and Atlantic Ocean (cosmopolitan). North-East Atlantic: first found in 1914 in La Rochelle (France) and 1937 in Sussex (UK). Scattered established populations present in some places around UK, Ireland and in some European countries, from The Netherlands to Portugal (Breton et al. 1995; Eno et al. 1997; Wolff 2005; Minchin 2007). Recorded from Mediterranean and Black Sea (de Kluijver and Ingalsuo 2009). Biology/ecology: Lives in warm waters (settlement may be aided by thermal effluents from power stations). Impacts: Competition with Semibalanus balanoides and Austrominius modestus (Kerckhof 2002). Vector(s): shipping, larvae in ballast waters, or fouling on ships’ hulls. Status: Introduced.

Amphibalanus eburneus Gould, 1841 (Fig. 1)

Fig. 1

Amphibalanus eburneus (Berre (France) © F. André)

The ivory barnacle. Native to Atlantic coast of Americas, from Boston to Rio de Janeiro. Introduced to many regions around the world including the Mediterranean and Black Sea and Pacific Ocean. North-East Atlantic: The Netherlands, France, Spain (Reise et al. 1999, Goulletquer et al. 2002, Wolff 2005). Biology/ecology: Often associated with mussels or other bivalves. Vector(s): fouling on ships’ hulls, ballast water, possibly mariculture. Status: Introduced.

Amphibalanus improvisus Darwin, 1854

The bay barnacle or acorn barnacle. Native to the North Atlantic Ocean. Introduced to Indo-Pacific, Australasia, Japan, Pacific North America, and elsewhere. North-East Atlantic: Established from Norway and The Netherlands to Spain, Ireland and UK (Reise et al. 1999; Wolff 2005; Minchin 2007). Biology/ecology: marine and brackish environments with low salinity (Polk 1963), ports, pontoons (Breton et al. 1995; Kerckhof 2002), sometimes as epibionts on crabs. Feeds on detritus and phytoplankton. Impacts: This species is regarded as a pest and is given as one of the 100 worst alien species in Europe (DAISIE 2009). Competes with other alien species such as Amphibalanus amphitrite and Austrominius modestus. Vector(s): Often dispersed by shipping: ballast water, fouling on hulls. Status: Cryptogenic; considered as introduced by some authors and native by others (see discussions in Kerckhof 2002; Wolff 2005; Kerckhof et al. 2007).

Amphibalanus reticulatus (Utinomi, 1978) [Balanus reticulatus].

Native to Indian Ocean, Indo-West Pacific, and invasive to Southwest Atlantic. North-East Atlantic: Belgium (Kerckhof 2002). Biology/ecology: fouling species, on buoys. Vector(s): ship fouling, ballast water. Status: Introduced.

Amphibalanus variegatus (Darwin, 1854)

Native to Indo-Malayan and Australia. North-East Atlantic: Reported from Belgium on buoys in 1997 and 1999 (Kerckhof 2002). Biology/ecology: on man-made structures, sheltered bays. Vector(s): ship fouling, ballast water. Status: Introduced.

Balanus trigonus Darwin, 1854 (Fig. 2)

Fig. 2

Balanus trigonus (Thau (France) 15/12/2006 © F. André)

Native to Pacific and Indian Oceans. North-East Atlantic: First reported from Azores in 1887 (Cardigos et al. 2006), now reported from The Netherlands and Belgium (Adema 1990; Kerckhof and Cattrijsse 2002). Biology/ecology: sublittoral, on floating objects, on invertebrates (Kerckhof 2002). Vector(s): ship fouling (Zullo 1992). Status: Introduced.

Megabalanus coccopoma (Darwin, 1854)

The titan acorn barnacle. Native to Pacific coasts of central and South America. North-East Atlantic: The Netherlands to France; established in Belgium in 1976 (Wolff 2005). Biology/ecology: Shallow waters, often found on buoys in ports. Vector(s): fouling on ships’ hulls and other man-made structures (Newman and McConnaughey 1987). Comments: First recorded in 1851, probably on a ship’s hull from Le Havre, France (Kerckhof and Cattrijsse 2002). Status: Introduced.

Megabalanus tintinnabulum (Linnaeus, 1758)

The giant barnacle. Native to tropical seas. North-East Atlantic: France, The Netherlands and Ireland, and established in Belgium (Wolff 2005; Minchin 2007). Biology/ecology: hard substrates in ports (hulls, buoys). Impacts: Unknown. Vector(s): fouling on ships’ hulls. Status: Cryptogenic.

Archeobalanidae

Solidobalanus fallax (Broch, 1927)

Native to African west coast. North-East Atlantic: Spain, Portugal, France and Great Britain (Eno et al. 1997; Kerckhof 2002). Biology/ecology: On lobster or crab pots, floating objects, shells such as the bivalve Aequipecten opercularis (Kerckhof, 2002). Vector(s): shipping, fouling. Status: Introduced.

Mysidacea

Mysidae

Hemimysis anomala G. O. Sars, 1907 (Fig. 3)

Fig. 3

Hemimysis anomala (Torcy (France) 16/05/2009 © J. Dumas)

The bloody-red mysid or Ponto-Caspian Mysid). Native to Ponto-Caspian area. North-East Atlantic: The Netherlands, Belgium, France, Ireland (Faasse 1998; Devin et al. 2005; Wolff 2005; Holdich and Pöckl 2007; Minchin and Holmes 2008). Biology/ecology: Fresh waters (rivers) and brackish waters (estuaries). Impacts: Dramatic effects on zooplankton composition and abundance (Ketelaars et al. 1999). Vector(s): shipping, ballast water (Ketelaars et al. 1999). Intentionally introduced for enrichment of fish feed in tributaries of the Baltic Sea (Salemaa and Hietalahti 1993). Status: Introduced.

Amphipoda

Caprellidae

Caprella mutica Schurin, 1935 [Caprella macho Platvoet, De Bruyne and Gmelich Meyling, 1995] (Fig. 4)

Fig. 4

Caprella mutica ((UK) © E. Cook)

The Japanese skeleton shrimp. Native to North-East Asia. Introduced to many regions in the world. North-East Atlantic: Celtic Sea, Ireland, England, Scotland, North Sea, Norway, Germany, The Netherlands (first record in Europe in 1994: Platvoet et al. 1995), Belgium, English Channel, and France (Breton 2005; Wolff 2005; Minchin 2007). Biology/ecology: Occurs in high numbers on artificial structures (boats hulls, buoys, pontoons, ropes and nets, aquaculture equipments) and biogenic reefs constructed by mussels (Modiolus modiolus, Mytilus edulis) and tubeworms (Sabellaria alveolata and Serpula vermicularis) (Cook et al. 2007a, b). Impacts: Largely unknown, but see Boos et al., this volume. Vector(s): ship hull fouling. For details on this species, see Boos et al. 2010. Status: Introduced.

Corophiidae

Chelicorophium curvispinum (Sars, 1895)

The Caspian mud shrimp. Native to Ponto-Caspian area. Also introduced to Canada and USA. North-East Atlantic: Ireland, England, The Netherlands, Germany, Belgium, France (Wundsch 1912; Devin et al. 2005; Wolff 2005; Holdich and Pöckl 2007; Minchin 2007). Biology/ecology: Salt, brackish and fresh waters. Rivers and estuaries (Buckley et al. 2004; de Kluijver and Ingalsuo 2009). Impacts: altering invaded habitats by predation, competition and causing changes to the substrate (Wittenberg 2006). Vector(s): Natural expansion through canals and rivers, facilitated by fouling on ships’ hulls. Status: Introduced.

Corophium multisetosum Stock, 1952

Native distribution undefined. Distributed along Atlantic coasts of Europe and Baltic; North-East Atlantic: Ireland, Wales, England, Germany, The Netherlands, France, Spain, Portugal (Stock 1952; Hayward and Ryland 1990; Bachelet et al. 2003). Biology/ecology: In brackish fish ponds (Bachelet et al. 2003); estuaries, near the limit of saline penetration (Buckley et al. 2004); burrows and constructs mud tubes, sometimes with seagrass Zostera noltii (de Kluijver and Ingalsuo 2009). Vector(s): Hull fouling. Comments: Usually considered as alien in different countries, but whether this species is introduced or not is questionable. The recent occurrence of C. multisetosum in German Baltic waters (Zettler et al. 2000) is possibly due to a natural range extension (Leppäkoski et al. 2003). Status: Cryptogenic.

Monocorophium acherusicum (da Costa, 1851 [nom. nud.] 1857) [Corophium acherusicum Costa, 1851 Corophium ascherusicum]

Native range uncertain; first described from Italy. North-East Atlantic: France, Belgium, England (Naylor 1965; Bachelet et al. 2003). Biology/ecology: constructs tubes on algae, hydroids and other organisms, in shallow sublittoral habitats of reduced salinity, often in sheltered harbours and estuaries. Vector(s): ballast water, ballast sediments or fouling on ships’ hulls. Status: Cryptogenic.

Monocorophium sextonae Crawford, 1937

The New Zealand mud shrimp. Native to New Zealand. North-East Atlantic: First introduced to Plymouth (UK) in 1930s (Crawford 1937), and later to Ireland. At present recorded from Ireland, Scotland, England and Germany to The Netherlands, Belgium, France, Portugal (Eno et al. 1997; Wolff 2005). Also present in Mediterranean (de Kluijver and Ingalsuo 2009). Biology/ecology: estuaries, open sea on Laminaria, Himanthalia, Buccinum eggs, hard substrates (Wolff 2005). Vector(s): ballast water or ballast sediments, fouling on ships’ hulls. Status: Introduced (Kerckhof et al. 2007).

Grandidierella japonica Stephensen, 1938

The Japanese amphipod. Native to China, Japan, Korea; introduced to Australia, Hawaii, Pacific coast of USA (Chapman and Dorman 1975). North-East Atlantic range: Suffolk (UK), North Sea (Smith et al. 1999; Ashelby 2006). Biology/ecology: tube dwelling species on sediments; brackish waters. Vector(s): ballast water, ballast sediments or fouling on ships’ hulls. Status: Introduced.

Gammaridae

Echinogammarus berilloni (Catta, 1871) [Gammarus berilloni]

An Iberian amphipod. Native to Atlantic part of Southwestern Europe: Spain, France (Pinkster 1973; Bachelet et al. 2003). North-East Atlantic: The Netherlands, Belgium, Luxembourg, Germany, Northern and eastern France (Peeters et al. 2003; Devin et al. 2005; Holdich and Pöckl 2007). Biology/ecology: Estuaries, brackish waters, canals, also inland waters. Vector(s): Floating structures, shipping (ballast water and/or sediments). Status: Introduced.

Gammarus tigrinus Sexton, 1939

The tiger sideswimmer or tiger gammarid. Native to Atlantic coast of North America, from St Lawrence Estuary, Canada, to Florida. North-East Atlantic: Ireland, Germany, The Netherlands, Belgium and France (Schmitz 1960; Devin et al. 2005; Wolff 2005; Minchin 2007). Common in the Baltic Sea. Biology/ecology: Estuaries, brackish waters, canals, also inland waters (de Kluijver and Ingalsuo 2009). Impacts: predatory impacts on macroinverebrates, can outcompete other amphipods (Pinkster et al. 1977). Vector(s): ballast water (Carlton 1985); deliberate introductions (Schmitz 1960; Bulnheim 1985). Status: Introduced.

Pleustidae

Incisocalliope aestuarius (Watling and Maurer, 1973) [Parapleustes assimilis (G. O. Sars, 1882); Pleusymtes glaber (Boeck, 1861)]

Native to Atlantic coast of North America (USA). North-East Atlantic: The Netherlands and Belgium (Faasse and Moorsel 2003, Wolff 2005). Biology/ecology: Estuaries (Kerckhof et al. 2007). Vector(s): ballast water, hull fouling. Status: Introduced.

Melitidae

Melita nitida Smith, 1837

Native to Atlantic coast of North America. North-East Atlantic: The Netherlands (Faasse and van Moorsel 2003, Wolff 2005). Biology/ecology: Estuaries, under stones, among sediments. Vector(s): ballast water or sediment, ships’ hull fouling. Status: Introduced.

Talitridae

Orchestia cavimana Heller, 1865

A freshwater riparian amphipod or semi-terrestrial amphipod. Native to Ponto-Caspian and Eastern-Mediterranean; Red Sea. North-East Atlantic: Germany, The Netherlands, Belgium, France, Spain (Tétry 1939; Kinzelbach 1972; Bachelet et al. 2003; Devin et al. 2005; Wolff 2005; Gollasch and Nehring 2006). Also along the Atlantic coast of Africa (de Kluijver and Ingalsuo 2009). Biology/ecology: demersal, semiterrestrial, under stones close to brackish waters, fresh waters. Vector(s): expansion along shipping canals (Kinzelbach 1972, 1995). Status: Introduced.

Platorchestia platensis (Krøyer, 1845) [Orchestia platensis]

The beach flea or sandhopper. Native distribution unknown, considered to be world-wide, but a likely species complex (Lowry 2000; Spicer and Janas 2006). North-East Atlantic: Specimens present or populations established in Norway, Sweden, The Netherlands, Germany (Persson 2001; Wolff 2005) [Baltic: Sweden, Denmark, Poland]. Biology/ecology: in wrack beds on hard substrates such as rocks, stones, gravels and shore meadows. Vector(s): Possibly dry ballast or natural dispersal with drifting algae. Status: Cryptogenic.

Tanaidacea

Tanaidae

Sinelobus stanfordi (Richardson, 1901) [Tanais stanfordi]

A brackish water tanaid. Native distribution unknown, reported as cosmopolitan (WoRMS 2010), and possibly a species complex; Pacific: Japan, New Zealand, and USA; Central West Atlantic: Mexico and USA (Vittor 2001). Central East Atlantic: Cameroon-Nigeria, South Africa. North-East Atlantic: Belgium, The Netherlands (van Haaren and Soors 2009). Biology/ecology: brackish waters of estuaries, ports. Vector(s): hull fouling, ballast water. Status: Cryptogenic.

Isopoda

Limnoriidae

Limnoria lignorum (Rathke, 1799) [Cymothoa lignora]

The common gribble. Native distribution still uncertain; a boreal species, recorded from the East and West coasts of North America (Schotte et al. 1995; de Kluijver and Ingalsuo 2009). North-East Atlantic: Reported from Scandinavia, UK, The Netherlands, France (Jones 1963; Wolff 2005). Biology/ecology: Lives in brackish waters of estuaries, ports. Impacts: reported as an important wood borer in many regions. Vector(s): Widely dispersed in the days of wooden sailing ships. Comments: Cryptogenic; see discussion in Wolff (2005).

Limnoria quadripunctata Holthuis, 1949

The quadripunctate gribble. Probably native to the South Pacific or Indian Oceans (Carlton, pers. comm. 2009), and introduced widely around the world (Schotte et al. 1995). North-East Atlantic: Described from The Netherlands, and reported from there, England, Ireland, France, and Spain (Jones 1963; Wolff 2005). Biology/ecology: Lives in brackish waters of estuaries, ports. Impacts: unknown in Europe, but reported as an important wood borer in many regions. Vector(s): Widely dispersed in the days of wooden sailing ships. Status: Introduced.

Limnoria tripunctata Menzies, 1951

The tripunctate gribble. Probably native to the South Pacific or Indian Oceans (Carlton and Eldredge 2009), and introduced widely around the world (Schotte et al. 1995). North-East Atlantic: British Isles (Jones 1963); also present in Mediterranean (Bourdillon 1958). Biology/ecology: wood boring isopod. Impacts: unknown in Europe, but reported as an important wood borer in many regions. Vector(s): Widely dispersed in the days of wooden sailing ships. Status: Introduced.

Idoteidae

Synidotea laticauda Benedict, 1897 [Synidotea laevidorsalis pro-parte]

Native to Japan, but species-level taxonomy remains in dispute (Chapman and Carlton 1991; 1994; Poore 1996). Also introduced to the Atlantic coast of the USA. North-East Atlantic: Reported from Gironde estuary, France since 1975 (Mees and Fockedey 1993 as S. laevidorsalis) and later from South-West Spain in 1996 (Cuesta et al. 1996 as S. laevidorsalis; Poore 1996; Drake et al. 2002 as S. laticauda). Biology/ecology: Littoral zone. Lives in brackish waters of estuaries and reported from docks, buoys and floating lines. Vector(s): Ship fouling or mariculture (oysters). Status: Introduced.

Decapoda

Portunidae

Callinectes sapidus Rathbun, 1896 (Fig. 5)

Fig. 5

Callinectes sapidus (Dunkerque (France) © P. Lesur)

The American blue crab. Native to Western Atlantic, from Canada to Argentina. Introduced to Japan, Pacific, as well as to Mediterranean (Galil et al. 2002). North-East Atlantic: Records are first from France (Bouvier 1901) then from Denmark, Germany, The Netherlands, Belgium, France, Spain and Portugal (d’Udekem d’Acoz 1999; Wolff 2005; Nehring 2010). Biology/ecology: brackish waters, estuaries, ports (Vincent 1986); reproduction is not really documented on North-East Atlantic. Impacts: Low in region under consideration, since records are scarce on North-East Atlantic. Vector(s): ballast water (larvae). Status: Introduced.

Majidae

Chionoecetes opilio (O. Fabricius, 1788) [Cancer opilio]

The snow crab. Naturally distributed in North Pacific and North-West Atlantic: Canada, Saint Pierre and Miquelon, USA, and Greenland (Tremblay 1997). North-East Atlantic: Barents Sea, Norway and Russia (d’Udekem d’Acoz 1999; Alvsvåg et al. 2009; Fey 2009). Biology/ecology: Lives in cold and deep waters. Impacts: Competition with other crabs. Vector(s): ballast water (larvae). For details on this species, see Agnalt et al. 2010. Status: Introduced.

Panopeidae

Dyspanopeus sayi (Smith, 1869) [Neopanope sayi]

The Say mud crab. Native to North-West Atlantic Ocean (USA). North-East Atlantic: introduced into Swansea Docks, Wales (Naylor 1960; d’Udekem d’Acoz 1999) and Mediterranean (Galil et al. 2002). Biology/ecology: estuaries and coastal lagoons. Impacts: Unknown for North-East Atlantic; elsehere may affect local clam farming (Galil et al. 2002). Vector(s): ballast water or mariculture. Status: Introduced.

Rhithropanopeus harrisii (Gould, 1841) [Pilumnus tridentatus Maitland, 1874; Heteropanope tridentate; Pilumnus harrisii] (Fig. 6)

Fig. 6

Rhithropanopeus harrisii (Veerse Meer (The Netherlands) © V. Maran)

The estuarine or Harris or white-tipped mud crab, Zuiderzee crab. Native to West-Atlantic from New Brunswick to North-East Brazil. North-East Atlantic: This crab is the first known decapod to be introduced to Europe, having first been collected in the 1870s in the Zuiderzee, where it was mistakenly described as a new species. It is established at present in most European countries (Eno et al. 1997). Population size has fluctuated recently; in some places, the crab has almost disappeared (Christiansen 1969; Ingle 1980; d’Udekem d’Acoz 1999; Wolff 2005). Also introduced into the Mediterranean (Galil et al. 2002). Biology/ecology: Lives in muddy waters of low salinity. Sometimes associated with the tube worm Ficopomatus enigmaticus. Impacts: Competes with native crabs (Marchand and Saudray 1971). Vector(s): Shipping. Status: Introduced.

Varunidae

Eriocheir sinensis H. Milne-Edwards, 1853 (Fig. 7)

Fig. 7

Eriocheir sinensis (Dunkerque (France) 27/04/2008 © F. Cordier)

The Chinese mitten crab. Native to North-West Pacific (Asia) between 40° N (Japan and Korean Peninsula) and 26° N (China). North-East Atlantic: First found in Germany in 1912, subsequently spread to other European countries, from Finland to Portugal. Populations of this crab increased in southern England, especially in Thames River, by the end of twentieth century. By contrast, the mitten crab is quite rare in France, for instance in Seine River (Vincent 1996). Biology/ecology: Catadromous; this crab is able to migrate long distances along rivers, and reproduces in brackish water. Impacts: It is regarded as a pest and is considered one of the 100 worst alien species in Europe (DAISIE 2009). It may impact soft sediment banks through burrowing. Vector(s): larvae in ballast water. Comments: Multiple invasions seemed to have occurred (Hänfling et al. 2002). For more details see Bentley (2010). Status: Introduced.

Hemigrapsus sanguineus (De Haan, 1835) (Fig. 8)

Fig. 8

Hemigrapsus sanguineus (Le Havre (France) 07/03/2009 © D. Ingratta)

The Asian or Japanese shore crab. Native to North-West Pacific (Asia), from Sakhalin Island to Taiwan. Also introduced to North-West Atlantic/USA, and to the Mediterranean (Galil et al. 2002). North-East Atlantic: France, The Netherlands (Breton et al. 2002; Campbell and Nijland 2004; Wolff 2005). Whether the European populations originate directly from Asia or indirectly from USA is unknown. Biology/ecology: Rocky places with algae, under stones. Impacts: Competition with other crabs such as Carcinus. Vector(s): ballast water, fouling. Status: Introduced.

Hemigrapsus takanoi Asakura and Watanabe, 2005 [Hemigrapsus penicillatus (De Haan, 1835) pro-parte: European specimens] (Fig. 9)

Fig. 9

Hemigrapsus tanakoi (Zeeland (The Netherlands) 05/04/2009 ©J.-P. Corolla)

The brush-clawed penicilate shore crab. Systematic note: Soon after the description of H. takanoi (Asakura and Watanabe 2005; Asakura 2006), Sakai (2007) synonymised it with Hemigrapsus penicillatus. After carefully examinating a large series of specimens, Ng et al. (2008) considered both species as valid. All European specimens so far examined are referrable to H. takanoi. Native to North-West Pacific (Asia), from Sakhalin Island to Taiwan. North-East Atlantic: Established first in France and Spain (Noël et al. 1997) then in The Netherlands, Belgium (Vincent and Breton 1999; Wolff 2005). Since 1995, populations developed swiftly in Charente-maritime, France then expanded north and south, reported from most coastal areas of Bay of Biscay (Noël et al. 1997; Noël and Gruet 2008). Biology/ecology: Omnivorous. Lives beneath stones and among empty oyster shells on mud flats; estuaries, lagoons, sheltered places. Impacts: Competition with other crabs such as Carcinus maenas and Pachygrapsus marmoratus. Southern expansion in Spain likely limited by competition with Eriphia spinifrons. Vector(s): Hull fouling, in empty barnacles (Gollasch 1999), ballast water or sediment, oyster mariculture. Six juveniles (of H. takanoi or H. penicillatus) were collected on the hull of a Japanese vessel in Bremerhaven, Germany, on 14 August 1993 (Gollasch 1999). Status: Introduced.

Palaemonidae

Palaemon macrodactylus Rathbun, 1902

The East Asian or oriental shrimp. Native to North-West Pacific (Asia). North-East Atlantic: in most large estuaries in UK, Germany, The Netherlands, Belgium, France and Spain (Cuesta et al. 2004; Ashelby et al. 2004; Béguer et al. 2007; Kerckhof et al. 2007; González-Ortegón et al. 2009). Biology/ecology: Large estuaries, marine and brackish waters. Impacts: Competition with other estuarine palaemonids. Vector(s): ballast water. Status: Introduced.

Lithodidae

Paralithodes camtschaticus (Tilesius, 1815) [Paralithodes camtschatica]

The Alaskan red king crab. Taxon: Lithodidae. Native to the North Pacific, Bering Sea, Japan Sea. North-East Atlantic range: Larvae, juveniles and adults introduced to the southern Russian Barents Sea (1961–1969), established and spread towards the north Norwegian coast (Finmark and South of the Lofoten Archipelago) by natural dispersion (Jørgensen and Primicerio 2007). Biology/ecology: Predator feeding on a large spectrum of benthic organisms. Impacts: The species is regarded as a pest and considered one of the 100 worst alien species in Europe (DAISIE 2009). Extensive interactions with marine macrofauna (Chlamys islandica) and algae (Gudimov et al. 2003). Vector(s): Deliberate introductions of larvae and adults. Comments: For details, see Anisimova et al. 2004; Jørgensen 2010. Status: Introduced.

Appendix B Species Here Considered to be Native or Non-established Aliens in Atlantic Europe

Cirripedia

Balanidae

Fistulobalanus albicostatus (Pilsbry, 1916)

A mangrove barnacle. Native to East Asia, Korea, Japan, Hong Kong, South China Sea, Taiwan. North-East Atlantic: France, some records from Bourgneuf Bay and Le Croisic (1974) but not established (Gruet and Baudet 1997; Goulletquer et al. 2002, 2004). Biology/ecology: Lives in warm waters, mangroves, estuaries, on ships, buoys, piers, docks. Vector(s): Mariculture.

Megabalanus tulipiformis (Ellis, 1758)

An acorn barnacle. Native to West coast of Africa; introduced to Mediterranean. North-East Atlantic: France, Spain and Madeira (Wirtz et al. (2006). Probably not established. Biology/ecology: hard substrates in ports (piers, buoys). Vector(s): fouling on ship’s hulls. Comments: Known as fossils from Miocene in France (Carriol 2004).

Archeobalanidae

Chirona hameri (Ascanius, 1767) [Balanus hameri]

An acorn barnacle. Native to North Atlantic deep water species. North-East Atlantic range: Reported from Belgium (Kerckhof 2002). Biology/ecology: Deep waters. Vector(s): fouling.

Lepadidae

Conchoderma auritum (Linnaeus, 1767)

The rabbit-ear barnacle. Cosmopolitan species: South Africa, Madagascar (Jones et al. 2000; WoRMS 2009). North-East Atlantic range: Faeroe Islands, Iceland, Norway, Sweden, Irish Sea, Scotland, North Sea, The Netherlands, Belgium, France [Wimereux], Madeira (Wolff 2005; WoRMS 2009; SeaLifeBase 2009). Biology/ecology: boat hulls, drift wood and attached to whale barnacles (de Kluijver and Ingalsuo 2009). Vector(s): Shipping, on ship’s hulls. Comments: Whether this species can be considered as introduced to Europe is not clear; According to DAISIE (2009) it is introduced; native according to SeaLifeBase.

Stomatopoda

Gonodactylidae

Odontodactylus scyllarus (Linnaeus, 1758)

A mantis shrimp. Indo-West Pacific. North-East Atlantic range: Saint Malo, Brittany (France) (Noël, unpublished data 2009). Biology/ecology: Impacts: weak if any. Vector(s): Possibly aquarium trade. Comments: A single specimen photographed in situ by a diver.

Decapoda

Varunidae

Brachynotus sexdentatus (Risso, 1827)

A grapsoid crab. Native to Mediterranean (references in d’ Udekem d’Acoz C 1999). North-East Atlantic: Introduced into Queens Dock, Swansea, Wales (Naylor 1957), then became extinct (Clark 1986). Further records are scarce, for instance in France at Roscoff [larvae] (Bourdon 1965) and La Rochelle [two specimens] (Noël et al. 1997). Biology/ecology: Lives in shallow waters, on sandy or rocky coasts in sheltered places. Vector(s): Ballast water, fouling, mariculture. There is also a possibility that this species is extending its natural geographical range further north due to climate warming.

Nephropidae

Homarus americanus H. Milne Edwards, 1837

The American lobster. Native to Atlantic coasts of North America, from Newfoundland to North-Carolina. North-East Atlantic range: Scattered records of specimens that are most likely escaped from captivity are available for Norway (Oslofjord), Sweeden, Danemark (Øresund), Ireland, UK, and France (d’Udekem d’Acoz 1999; Minchin 2007); it is not known if reproducing populations are established. Biology/ecology: Rocky shores, infralittoral. Vector(s): Mariculture, imports of live crustaceans for human consumption. For details, see Jørstad et al. 2010.

Palinuridae

Jasus lalandii (H. Milne Edwards, 1837)

The Cape rock lobster. Native to South Africa, Namibia. North-East Atlantic: Portugal (Guerra and Gaudêncio 1982), no further records; Biology/ecology: Marine, rocky bottoms. Vector(s): Intentional release.

Penaeidae

Marsupenaeus japonicus (Bate, 1888) [Penaeus japonicus]

The Kuruma shrimp. Native throughout Indo-Pacific including Red Sea. Introduced into Mediterranean (Galil et al. 2002). North-East Atlantic: Some records are available for specimens escaped from aquaculture facilities (Clark 1990a, b; d’Udekem d’Acoz 1999; Minchin 2007). Biology/ecology: Lives in coastal waters; reproduction is not documented on North-East Atlantic coast. Impacts: This shrimp is regarded as a pest and is listed as one of the 100 worst alien species in Europe (DAISIE 2009). Vector(s): Mariculture.

Menippidae

Menippe mercenaria (Say, 1818)

The Florida stone crab. Native to Atlantic coasts of USA, Mexico, Cuba. North-East Atlantic range: One living adult female present among fouling in Brittany, France on a buoy originating from Florida and drifting for 18 month across North Atlantic (Noël 2007). Biology/ecology: Tolerant to various salinities and preys on oysters and other molluscs. Burrows in mud and also present on hard bottoms (Tavares 2002). Vector(s): Intercontinental drift on floating objects.

Pilumnoididae

Pilumnoides inglei Guinot and Macpherson, 1987

Native distribution: Probably South America; Pilumnoides has tropical representatives only. North-East Atlantic range: Early 1900s records from Ireland and England; not recorded since 1913 and considered not established (Vallentin 1900; Ingle 1980; Clark 1986; Guinot and Macpherson 1987; d’Udekem d’Acoz 1999). Biology/ecology: From the hulls of ships docking in ports. Vector(s): Fouling. Comments: Carlton (2009) reviewed the history of this species, which was described as new from the British Isles: however, Guinot and MacPherson (1987) could find no significant differences between it and the South American P. perlatus (Poeppig, 1836). Ng et al. (2008) retain it as a distinct species.