CARCINUS MAENAS
(Redirected from European Green Crab)
'''Carcinus maenas''' is a common littoral crab, and an important invasive species. It is listed among the 100 "world's worst invasive alien species" [1].
''C. maenas'' is known by different names around the world. In the British Isles, it is generally referred to simply as the ''shore crab''. In North America and South Africa, it bears the name ''green crab'' or ''European green crab''. In Australia and New Zealand, it is referred to as either the ''European green crab'' or ''European shore crab''.
''C. maenas'' has a carapace up to 60 mm long and 90 mm wide [2], with five short teeth along the rim behind each eye, and three undulations between the eyes. The undulations, which do not protrude beyond the eyes are the simplest means of distinguishing ''C. maenas'' from the closely-related ''C. aestuarii'', which can also be an invasive species. In ''C. aestuarii'', the carapace lacks any bumps and extends forward beyond the eyes. The other character for distinguishing the two species is the form of the first and second pleopods (collectively the gonopods), which are straight and parallel in ''C. aestuarii'', but curve outwards in ''C. maenas'' .
The colour of ''C. maenas'' varies greatly, from green to brown, grey or red. This variation has a genetic component but is largely due to local environmental factors [3]. In particular, individuals which delay moulting become red–coloured rather than green. Red individuals are stronger and more aggressive, but are less tolerant of environmental stresses, such as low salinity or hypoxia [4].
''C. maenas'' is native to European and North African coasts as far as the Baltic Sea in the east, and Iceland and central Norway in the north, and is one of the most common crabs throughout much of its range. In the Mediterranean Sea, it is replaced by the closely-related species ''Carcinus aestuarii''.
''C. maenas'' was first observed on the east coast of North America in Massachusetts in 1817, and may now be found from Nova Scotia to Virginia.
In Australia, ''C. maenas'' was first reported in the late 19th century, in Port Phillip Bay, Victoria. It has since spread along the coast of Victoria, reaching New South Wales in 1971, South Australia in 1976 and Tasmania in 1993. One specimen was found in Western Australia in 1965, but ''C. maenas'' has not been seen in the area since [5].
''C. maenas'' first reached South Africa in 1983, in the Table Docks area near Cape Town [6]. Since then, it has spread at least as far as Saldanha Bay in the north and Camps Bay in the south, over 100 km apart.
In 1989, ''C. maenas'' was found in San Francisco Bay, California, on the Pacific coast of the United States. Until 1993, it was not able to extend its range, but reached Oregon in 1997, the state of Washington in 1998 and British Columbia in 1999 [7][8].
In 2003, ''C. maenas'' was discovered in Patagonia [9].
Other appearances that have not, however, led to invasions have been recorded in Brazil, Panama, Hawaii, Madagascar, the Red Sea, Pakistan, Sri Lanka and Myanmar; Japan has been invaded by a related crab, either ''C. aestuarii'' or a hybrid of ''C. aestuarii'' and ''C. maenas'' [10].
It is believed, based on the ecological conditions, that ''C. maenas'' could eventually extend its range to colonise the Pacific coast of North America from Baja California to Alaska 7. Similar ecological conditions are to be found on many of the world's coasts, with the only large potential area not to have been invaded yet being New Zealand; the New Zealand government has taken action, including the release of a Marine Pest Guide [11] in an effort to prevent colonisation by ''C. maenas''.
''C. maenas'' can live in all types of protected and semi-protected marine and estuarine habitats, including habitats with mud, sand, or rock substrates, submerged aquatic vegetation, and emergent marsh, although soft bottoms are preferred. ''C. maenas'' is euryhaline, meaning that it can tolerate a wide range of salinities (from 4 to 52 ‰)and survive in temperatures from 0°C to 30°C [12]. The wide salinity range allows ''C. maenas'' to survive in the lower salinities found in estuaries.
A molecular biological study using the COI gene found genetic differentiation between the North Sea and the Bay of Biscay, and even more strongly between the populations in Iceland and the Faroe Islands and those elsewhere. This suggests that ''C. maenas'' is unable to cross deeper water [13].
Females can produce up to 185,000 eggs and larvae develop offshore in several stages before their final moult to juvenile crabs in the intertidal zone USGS Nonindigenous Aquatic Species Database Perry . Young crabs live in meadows of kelp, for instance ''Posidonia oceanica'', until they reach adulthood [14].
''C. maenas'' is a predator and feeds on many organisms, particularly bivalve molluscs (such as clams, oysters, and mussels), polychaetes and small crustaceans [15]. They are primarily nocturnal, although activity also depends on the tide, and crabs can be active at any time of day [16].
''C. maenas'' has the ability to disperse by a variety of mechanisms including: ballast water, ships' hulls, packing materials (seaweeds) used to ship live marine organisms, bivalves moved for aquaculture, rafting, migration of crab larvae on ocean currents, and the movement of submerged aquatic vegetation for coastal zone management initiatives. Thresher ''et al.'' 5 found that in Australia ''C. maenas'' dispersed mainly by rare long-distance events, possibly caused by human actions.
In California, preferential predation of ''C. maenas'' on native clams (''Nutricola spp.'') resulted in the decline of the native clams and an increase of a previously introduced clam (the amethyst gem clam, ''Gemma gemma'') [17]. ''C. maenas'' has been implicated in the destruction of the soft-shell clam (''Mya arenaria'') fisheries on the east coast of the United States and Canada and the reduction of populations of other commercially important bivalves (such as scallops, ''Argopecten irradians'', and northern quahogs, ''Mercenaria mercenaria'') . The prey of ''C. maenas'' includes the young of bivalves [18] and fish, although the effect of its predation on winter flounder, ''Pseudopleuronectes americanus'' is minimal [19]. ''C. maenas'' can, however, have substantial negative impacts on local commercial and recreational fisheries by preying on the young of species such as oysters and the Dungeness crab, or competing with them for resources [20].
Due to its potentially harmful effects on ecosystems, various efforts have been made to control introduced populations of ''C. maenas'' around the world. In Edgartown, Massachusetts, a bounty was levied in 1995 for catching ''C. maenas'', in order to protect local shellfish, and 10 tonnes were caught [21].
There is evidence that the native blue crab in eastern North America, ''Callinectes sapidus'', is able to control populations of ''C. maenas''; numbers of the two species are negatively correlated, and ''C. maenas'' is not found in Chesapeake Bay, where ''Callinectes sapidus'' is most frequent [22]. On the west coast of North America, ''C. maenas'' appears to be limited to upper estuarine habitats in part by predation by native rock crabs (''Cancer antennarius'' and ''Cancer productus'') and competition for shelter with a native shore crab, ''Hemigrapsus oregonensis'' [23]. Host specificity testing has recently been conducted on ''Sacculina carcini'', a parasitic barnacle, as a potential biological control agent of ''C. maenas'' [24]. In the laboratory, ''Sacculina'' settled on, infected, and killed native California crabs, including the Dungeness crab, ''Cancer magister'', and the shore crabs ''Hemigrapsus nudus'', ''Hemigrapsus oregonensis'' and ''Pachygrapsus crassipes''. Dungeness crabs were the most vulnerable of the tested native species to settlement and infection by the parasite. Although ''Sacculina'' did not mature in any of the native crabs, developing reproductive sacs were observed inside a few ''Cancer magister'' and ''Hemigrapsus oregonensis''. Any potential benefits of using ''Sacculina'' to control ''C. maenas'' on the west coast of North America would need to be weighed against these potential non-target impacts .
''C. maenas'' has also caused significant problems in the Republic of Ireland where predation of bottom-cultured mussels has been recorded in several counties (Kerry, Wexford, Waterford), and a number of control programmes have been instigated by mussel producers. "Hard-eye" shrimp creels are typically used to fish ''C. maenas'' in Ireland; the creels are baited, deployed and fished in 24-hour cycles, being typically deployed in strings along the boundaries of mussel beds in order to lure the crabs out of the mussel beds. Some producers pay levies for the removal of crabs, but in most cases, ''C. maenas'' catches are sold for processing into food products. This is, however, a small market and the relatively low value of ''C. maenas'' (€500 per tonne) and high transport costs (refrigerated lorry €170 per tonne) means that fishing ''C. maenas'' is uneconomical unless it forms part of a control programme to protect more valuable shellfish beds .
''C. maenas'' is fished on a small scale in the north-east Atlantic Ocean, with approximately 1200 tonnes being caught annually, mostly in France and the United Kingdom. In the north-west Atlantic, ''C. maenas'' was the subject of fishery in the 1960s, and again since 1996, with up to 86 tonnes being caught annually [25].
1. 100 of the World's Worst Invasive Alien Species
2. Field identification of the European green crab species: ''Carcinus maenas'' and ''Carcinus aestuarii'', S. B. Yamada & L. Hauck, , , Journal of Shellfish Research, 2001
3. {{cite journal | quotes=no|title=Patterns of morphological and genetic variability in UK populations of the shore crab, ''Carcinus maenas'' Linnaeus, 1758 (Crustacea : Decapoda : Brachyura) |author=J. V. Brian, T. Fernandes, R. J. Ladle & P. A. Todd |year=2005 |journal=Journal of Experimental Marine Biology |volume=329 |issue=1 |pages=47–54 |id=
4. Distribution is correlated with color phase in green crabs, ''Carcinus maenas'' (Linnaeus, 1758) in southern New England, A. McKnight, L. M. Mathres, R. Avery & K. T. Lee, , , Crustaceana, 2000
5. Invasion dynamics of the European shore crab, ''Carcinus maenas'', in Australia, R. Thresher, C. Proctor, G. Ruiz, R. Gurney, C. MacKinnon, W. Walton, L. Rodriguez & N. Bax, , , Marine Biology, 2003
6. On the distribution, diet, and possible impact of the invasive European shore crab ''Carcinus maenas'' (L.) along the South African coast, P. J. Le Roux, G. M. Branch & M. A. P. Joska, , , South African Journal of Marine Science, 1990
7. Non-indigenous aquatic species of concern for Alaska. Fact Sheet 1
8. {{cite journal | quotes=no|author=E. D. Grosholz & G. M. Ruiz|year=1996|title=Predicting the impact of introduced marine species: Lessons from the multiple invasions of the European green crab ''Carcinus maenas''|journal=Biological Conservation |volume=78|issue=1-2|pages=59–66 |id=
9. {{cite journal | quotes=no|author=F. J. Hidalgo ''et al.''|year=2005|title=A prediction come true: the green crab invades the Patagonian coast |journal=Biological Invasions |volume=7 |issue=3|pages=547–552 |id=
10. Episodic global dispersal in shallow water marine organisms: the case history of the European shore crabs ''Carcinus maenas'' and ''C. aestuarii'', T. J. Carlton & A. N. Cohen, , , Journal of Biogeography, 2003
11. Marine Pest Guide
12. Introduction, dispersal and potential impacts of the green crab ''Carcinus maenas'' in San Francisco Bay, California, A. N. Cohen & T. J. Carlton, , , Marine Biology, 1995
13. A global invader at home: population structure of the green crab, ''Carcinus maenas'', in Europe, J. Roman & S. R. Palumbi, , , Molecular Ecology, 2004
14. Colour change and mimicry from juvenile to adult: ''Xantho poressa'' (Olivi, 1792) (Brachyura, Xanthidae) and ''Carcinus maenas'' (Linnaeus, 1758) (Brachyura, Portunidae), B. Bedini, , , Crustaceana, 2002
15. European green crab Washington Department of Fish and Wildlife
16. Diurnal activity in a group of Maine decapods, M. Novak, , , Crustaceana, 2004
17. Recent biological invasion may hasten invasional meltdown by accelerating historical introductions, E. D. Grosholz, , , PNAS, 2005
18. Predation potential of the invasive green crab (''Carcinus maenas'') and other common predators on commercial bivalve species found on Prince Edward island, G. Miron ''et al.'', , , Journal of Shellfish Research, 2005
19. {{cite journal | quotes=no|author=D. L. Taylor |year=2005|title=Predatory impact of the green crab (''Carcinus maenas'' Linnaeus) on post-settlement winter flounder (''Pseudopleuronectes americanus'' Walbaum) as revealed by immunological dietary analysis|journal=Journal of Experimental Marine Biology and Ecology |volume=324|issue=2|pages=112–126 |id=
20. Biological control of marine pests, K. D. Lafferty & K. D. Kuris, , , Ecology, 1996
21. The Web-Based Rapid Response Toolbox. Web publication F. R. McEnnulty, T. E. Jones & N. J. Bax
22. Biotic resistance to invasion: Native predator limits abundance and distribution of an introduced crab, DeRivera, C E, G M Ruiz, A H Hines & P Jivoff, , , Ecology, 2005
23. Global invader: the European Green Crab, S. B. Yamada, , , Oregon Sea Grant, 2001,
24. Host specificity of ''Sacculina carcini'', a potential biological control agent of introduced European green crab ''Carcinus maenas'' in California, J. H. R. Goddard, M. E. Torchin, A. M. Kuris & K. D. Lafferty, , , Biological Invasions, 2005
25. FAO Nominal Catches: ''Carcinus maenas''
'''Carcinus maenas''' is a common littoral crab, and an important invasive species. It is listed among the 100 "world's worst invasive alien species" [1].
''C. maenas'' is known by different names around the world. In the British Isles, it is generally referred to simply as the ''shore crab''. In North America and South Africa, it bears the name ''green crab'' or ''European green crab''. In Australia and New Zealand, it is referred to as either the ''European green crab'' or ''European shore crab''.
| Contents |
| Description |
| Native and introduced range |
| Ecology |
| Control |
| Fishery |
| References |
Description
''C. maenas'' has a carapace up to 60 mm long and 90 mm wide [2], with five short teeth along the rim behind each eye, and three undulations between the eyes. The undulations, which do not protrude beyond the eyes are the simplest means of distinguishing ''C. maenas'' from the closely-related ''C. aestuarii'', which can also be an invasive species. In ''C. aestuarii'', the carapace lacks any bumps and extends forward beyond the eyes. The other character for distinguishing the two species is the form of the first and second pleopods (collectively the gonopods), which are straight and parallel in ''C. aestuarii'', but curve outwards in ''C. maenas'' .
The colour of ''C. maenas'' varies greatly, from green to brown, grey or red. This variation has a genetic component but is largely due to local environmental factors [3]. In particular, individuals which delay moulting become red–coloured rather than green. Red individuals are stronger and more aggressive, but are less tolerant of environmental stresses, such as low salinity or hypoxia [4].
Native and introduced range
''C. maenas'' is native to European and North African coasts as far as the Baltic Sea in the east, and Iceland and central Norway in the north, and is one of the most common crabs throughout much of its range. In the Mediterranean Sea, it is replaced by the closely-related species ''Carcinus aestuarii''.
''C. maenas'' was first observed on the east coast of North America in Massachusetts in 1817, and may now be found from Nova Scotia to Virginia.
In Australia, ''C. maenas'' was first reported in the late 19th century, in Port Phillip Bay, Victoria. It has since spread along the coast of Victoria, reaching New South Wales in 1971, South Australia in 1976 and Tasmania in 1993. One specimen was found in Western Australia in 1965, but ''C. maenas'' has not been seen in the area since [5].
''C. maenas'' first reached South Africa in 1983, in the Table Docks area near Cape Town [6]. Since then, it has spread at least as far as Saldanha Bay in the north and Camps Bay in the south, over 100 km apart.
In 1989, ''C. maenas'' was found in San Francisco Bay, California, on the Pacific coast of the United States. Until 1993, it was not able to extend its range, but reached Oregon in 1997, the state of Washington in 1998 and British Columbia in 1999 [7][8].
In 2003, ''C. maenas'' was discovered in Patagonia [9].
Other appearances that have not, however, led to invasions have been recorded in Brazil, Panama, Hawaii, Madagascar, the Red Sea, Pakistan, Sri Lanka and Myanmar; Japan has been invaded by a related crab, either ''C. aestuarii'' or a hybrid of ''C. aestuarii'' and ''C. maenas'' [10].
It is believed, based on the ecological conditions, that ''C. maenas'' could eventually extend its range to colonise the Pacific coast of North America from Baja California to Alaska 7. Similar ecological conditions are to be found on many of the world's coasts, with the only large potential area not to have been invaded yet being New Zealand; the New Zealand government has taken action, including the release of a Marine Pest Guide [11] in an effort to prevent colonisation by ''C. maenas''.
Ecology
''C. maenas'' can live in all types of protected and semi-protected marine and estuarine habitats, including habitats with mud, sand, or rock substrates, submerged aquatic vegetation, and emergent marsh, although soft bottoms are preferred. ''C. maenas'' is euryhaline, meaning that it can tolerate a wide range of salinities (from 4 to 52 ‰)and survive in temperatures from 0°C to 30°C [12]. The wide salinity range allows ''C. maenas'' to survive in the lower salinities found in estuaries.
A molecular biological study using the COI gene found genetic differentiation between the North Sea and the Bay of Biscay, and even more strongly between the populations in Iceland and the Faroe Islands and those elsewhere. This suggests that ''C. maenas'' is unable to cross deeper water [13].
Females can produce up to 185,000 eggs and larvae develop offshore in several stages before their final moult to juvenile crabs in the intertidal zone USGS Nonindigenous Aquatic Species Database Perry . Young crabs live in meadows of kelp, for instance ''Posidonia oceanica'', until they reach adulthood [14].
''C. maenas'' is a predator and feeds on many organisms, particularly bivalve molluscs (such as clams, oysters, and mussels), polychaetes and small crustaceans [15]. They are primarily nocturnal, although activity also depends on the tide, and crabs can be active at any time of day [16].
''C. maenas'' has the ability to disperse by a variety of mechanisms including: ballast water, ships' hulls, packing materials (seaweeds) used to ship live marine organisms, bivalves moved for aquaculture, rafting, migration of crab larvae on ocean currents, and the movement of submerged aquatic vegetation for coastal zone management initiatives. Thresher ''et al.'' 5 found that in Australia ''C. maenas'' dispersed mainly by rare long-distance events, possibly caused by human actions.
In California, preferential predation of ''C. maenas'' on native clams (''Nutricola spp.'') resulted in the decline of the native clams and an increase of a previously introduced clam (the amethyst gem clam, ''Gemma gemma'') [17]. ''C. maenas'' has been implicated in the destruction of the soft-shell clam (''Mya arenaria'') fisheries on the east coast of the United States and Canada and the reduction of populations of other commercially important bivalves (such as scallops, ''Argopecten irradians'', and northern quahogs, ''Mercenaria mercenaria'') . The prey of ''C. maenas'' includes the young of bivalves [18] and fish, although the effect of its predation on winter flounder, ''Pseudopleuronectes americanus'' is minimal [19]. ''C. maenas'' can, however, have substantial negative impacts on local commercial and recreational fisheries by preying on the young of species such as oysters and the Dungeness crab, or competing with them for resources [20].
Control
Due to its potentially harmful effects on ecosystems, various efforts have been made to control introduced populations of ''C. maenas'' around the world. In Edgartown, Massachusetts, a bounty was levied in 1995 for catching ''C. maenas'', in order to protect local shellfish, and 10 tonnes were caught [21].
There is evidence that the native blue crab in eastern North America, ''Callinectes sapidus'', is able to control populations of ''C. maenas''; numbers of the two species are negatively correlated, and ''C. maenas'' is not found in Chesapeake Bay, where ''Callinectes sapidus'' is most frequent [22]. On the west coast of North America, ''C. maenas'' appears to be limited to upper estuarine habitats in part by predation by native rock crabs (''Cancer antennarius'' and ''Cancer productus'') and competition for shelter with a native shore crab, ''Hemigrapsus oregonensis'' [23]. Host specificity testing has recently been conducted on ''Sacculina carcini'', a parasitic barnacle, as a potential biological control agent of ''C. maenas'' [24]. In the laboratory, ''Sacculina'' settled on, infected, and killed native California crabs, including the Dungeness crab, ''Cancer magister'', and the shore crabs ''Hemigrapsus nudus'', ''Hemigrapsus oregonensis'' and ''Pachygrapsus crassipes''. Dungeness crabs were the most vulnerable of the tested native species to settlement and infection by the parasite. Although ''Sacculina'' did not mature in any of the native crabs, developing reproductive sacs were observed inside a few ''Cancer magister'' and ''Hemigrapsus oregonensis''. Any potential benefits of using ''Sacculina'' to control ''C. maenas'' on the west coast of North America would need to be weighed against these potential non-target impacts .
''C. maenas'' has also caused significant problems in the Republic of Ireland where predation of bottom-cultured mussels has been recorded in several counties (Kerry, Wexford, Waterford), and a number of control programmes have been instigated by mussel producers. "Hard-eye" shrimp creels are typically used to fish ''C. maenas'' in Ireland; the creels are baited, deployed and fished in 24-hour cycles, being typically deployed in strings along the boundaries of mussel beds in order to lure the crabs out of the mussel beds. Some producers pay levies for the removal of crabs, but in most cases, ''C. maenas'' catches are sold for processing into food products. This is, however, a small market and the relatively low value of ''C. maenas'' (€500 per tonne) and high transport costs (refrigerated lorry €170 per tonne) means that fishing ''C. maenas'' is uneconomical unless it forms part of a control programme to protect more valuable shellfish beds .
Fishery
''C. maenas'' is fished on a small scale in the north-east Atlantic Ocean, with approximately 1200 tonnes being caught annually, mostly in France and the United Kingdom. In the north-west Atlantic, ''C. maenas'' was the subject of fishery in the 1960s, and again since 1996, with up to 86 tonnes being caught annually [25].
References
1. 100 of the World's Worst Invasive Alien Species
2. Field identification of the European green crab species: ''Carcinus maenas'' and ''Carcinus aestuarii'', S. B. Yamada & L. Hauck, , , Journal of Shellfish Research, 2001
3. {{cite journal | quotes=no|title=Patterns of morphological and genetic variability in UK populations of the shore crab, ''Carcinus maenas'' Linnaeus, 1758 (Crustacea : Decapoda : Brachyura) |author=J. V. Brian, T. Fernandes, R. J. Ladle & P. A. Todd |year=2005 |journal=Journal of Experimental Marine Biology |volume=329 |issue=1 |pages=47–54 |id=
4. Distribution is correlated with color phase in green crabs, ''Carcinus maenas'' (Linnaeus, 1758) in southern New England, A. McKnight, L. M. Mathres, R. Avery & K. T. Lee, , , Crustaceana, 2000
5. Invasion dynamics of the European shore crab, ''Carcinus maenas'', in Australia, R. Thresher, C. Proctor, G. Ruiz, R. Gurney, C. MacKinnon, W. Walton, L. Rodriguez & N. Bax, , , Marine Biology, 2003
6. On the distribution, diet, and possible impact of the invasive European shore crab ''Carcinus maenas'' (L.) along the South African coast, P. J. Le Roux, G. M. Branch & M. A. P. Joska, , , South African Journal of Marine Science, 1990
7. Non-indigenous aquatic species of concern for Alaska. Fact Sheet 1
8. {{cite journal | quotes=no|author=E. D. Grosholz & G. M. Ruiz|year=1996|title=Predicting the impact of introduced marine species: Lessons from the multiple invasions of the European green crab ''Carcinus maenas''|journal=Biological Conservation |volume=78|issue=1-2|pages=59–66 |id=
9. {{cite journal | quotes=no|author=F. J. Hidalgo ''et al.''|year=2005|title=A prediction come true: the green crab invades the Patagonian coast |journal=Biological Invasions |volume=7 |issue=3|pages=547–552 |id=
10. Episodic global dispersal in shallow water marine organisms: the case history of the European shore crabs ''Carcinus maenas'' and ''C. aestuarii'', T. J. Carlton & A. N. Cohen, , , Journal of Biogeography, 2003
11. Marine Pest Guide
12. Introduction, dispersal and potential impacts of the green crab ''Carcinus maenas'' in San Francisco Bay, California, A. N. Cohen & T. J. Carlton, , , Marine Biology, 1995
13. A global invader at home: population structure of the green crab, ''Carcinus maenas'', in Europe, J. Roman & S. R. Palumbi, , , Molecular Ecology, 2004
14. Colour change and mimicry from juvenile to adult: ''Xantho poressa'' (Olivi, 1792) (Brachyura, Xanthidae) and ''Carcinus maenas'' (Linnaeus, 1758) (Brachyura, Portunidae), B. Bedini, , , Crustaceana, 2002
15. European green crab Washington Department of Fish and Wildlife
16. Diurnal activity in a group of Maine decapods, M. Novak, , , Crustaceana, 2004
17. Recent biological invasion may hasten invasional meltdown by accelerating historical introductions, E. D. Grosholz, , , PNAS, 2005
18. Predation potential of the invasive green crab (''Carcinus maenas'') and other common predators on commercial bivalve species found on Prince Edward island, G. Miron ''et al.'', , , Journal of Shellfish Research, 2005
19. {{cite journal | quotes=no|author=D. L. Taylor |year=2005|title=Predatory impact of the green crab (''Carcinus maenas'' Linnaeus) on post-settlement winter flounder (''Pseudopleuronectes americanus'' Walbaum) as revealed by immunological dietary analysis|journal=Journal of Experimental Marine Biology and Ecology |volume=324|issue=2|pages=112–126 |id=
20. Biological control of marine pests, K. D. Lafferty & K. D. Kuris, , , Ecology, 1996
21. The Web-Based Rapid Response Toolbox. Web publication F. R. McEnnulty, T. E. Jones & N. J. Bax
22. Biotic resistance to invasion: Native predator limits abundance and distribution of an introduced crab, DeRivera, C E, G M Ruiz, A H Hines & P Jivoff, , , Ecology, 2005
23. Global invader: the European Green Crab, S. B. Yamada, , , Oregon Sea Grant, 2001,
24. Host specificity of ''Sacculina carcini'', a potential biological control agent of introduced European green crab ''Carcinus maenas'' in California, J. H. R. Goddard, M. E. Torchin, A. M. Kuris & K. D. Lafferty, , , Biological Invasions, 2005
25. FAO Nominal Catches: ''Carcinus maenas''
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