Carcinus maenas

"Shore crab" redirects here. This may also be used for crabs in the superfamily Grapsoidea.

Carcinus maenas
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Crustacea
Class:	Malacostraca
Order:	Decapoda
Infraorder:	Brachyura
Family:	Portunidae
Genus:	Carcinus
Species:	C. maenas
Binomial name
Carcinus maenas (Linnaeus, 1758) [1]

Carcinus maenas is a common littoral crab, and an important invasive species, listed among the 100 "world's worst alien invasive species".^[2] It is native to the north-east Atlantic Ocean and Baltic Sea, but has colonised similar habitats in Australia, South Africa, South America and both Atlantic and Pacific coasts of North America. It grows to a carapace width of 90 millimetres (3.5 in), and feeds on a variety of molluscs, worms and small crustaceans, potentially impacting a number of fisheries. Its successful dispersion has occurred via a variety of mechanisms, such as on ships' hulls, packing materials, bivalves moved for aquaculture, and rafting.

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.

Description

A young Carcinus maenas showing the common green colour

C. maenas has a carapace up to 60 millimetres (2.4 in) long and 90 mm (3.5 in) wide,^[3] with five short teeth along the rim behind each eye, and three undulations between the eyes. The undulations, which 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. Another characteristic 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.^[3]

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.^[4] 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.^[5]

Native and introduced range

Rough map of the distribution of Carcinus maenas, blue areas are the native range, red areas are the introduced or invasive range, black dots represent single sightings that did not lead to invasion, and green areas are the potential range of the species.

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 southern Virginia northwards; by 2007, this species had extended its range northwards to Placentia Bay, Newfoundland.^[6] In 1989, the species 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] thus extending its range by 750 kilometres (470 mi) in ten years.^[9] By 2003, C. maenas had extended to South America with specimens discovered in Patagonia.^[10]

In Australia, C. maenas was first reported "in the late 1800s",^[11] in Port Phillip Bay, Victoria. It has since spread along the south-eastern and south-western seaboards, reaching New South Wales in 1971, South Australia in 1976 and Tasmania in 1993. One specimen was found in Western Australia in 1965, but there have been no further discoveries in the area since.^[11]

C. maenas first reached South Africa in 1983, in the Table Docks area near Cape Town.^[12] Since then, it has spread at least as far as Saldanha Bay in the north and Camps Bay in the south, over 100 kilometres (62 mi) apart.

There have been appearances of C. maenas recorded in Brazil, Panama, Hawaii, Madagascar, the Red Sea, Pakistan, Sri Lanka and Myanmar; however, these have not resulted in invasions, but remain isolated findings. Japan has been invaded by a related crab, either C. aestuarii or a hybrid of C. aestuarii and C. maenas.^[13]

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^[14] in an effort to prevent colonisation by C. maenas.

Ecology

A female Carcinus maenas carrying fertilised eggs

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 of 0 to 30 °C (32 to 86 °F).^[15] 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.^[16]

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.^[17] Young crabs live among seaweeds and seagrasses, such as Posidonia oceanica, until they reach adulthood.^[18]

Argopecten irradians, a scallop which has been affected by the introduction of C. maenas

C. maenas has the ability to disperse by a variety of mechanisms,^[17] 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.^[11] found C. maenas dispersed in Australia mainly by rare long-distance events, possibly caused by human actions.

C. maenas is a predator, feeding on many organisms, particularly bivalve molluscs (such as clams, oysters, and mussels), polychaetes and small crustaceans.^[19] They are primarily nocturnal, although activity also depends on the tide, and crabs can be active at any time of day.^[20] 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).^[21] 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).^[17] The prey of C. maenas includes the young of bivalves^[22] and fish, although the effect of its predation on winter flounder, Pseudopleuronectes americanus is minimal.^[23] 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.^[24]

Control

Cancer productus limits the spread of Carcinus maenas in parts of North America.

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, to protect local shellfish, and 10 tons were caught.^[25]

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 the Chesapeake Bay, where Callinectes sapidus is most frequent.^[26] On the west coast of North America, C. maenas appears to be limited to upper estuarine habitats, in part because of predation by native rock crabs (Romaleon antennarium and Cancer productus) and competition for shelter with a native shore crab, Hemigrapsus oregonensis.^[27] Host specificity testing has recently been conducted on Sacculina carcini, a parasitic barnacle, as a potential biological control agent of C. maenas.^[28] In the laboratory, Sacculina settled on, infected, and killed native California crabs, including the Dungeness crab, Metacarcinus magister (formerly 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 Metacarcinus 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.^[28]

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 northwest Atlantic, C. maenas was the subject of fishery in the 1960s, and again since 1996, with up to 86 tonnes being caught annually.^[29]

Taxonomic history

Carcinus maenas was first given a binomial name, Cancer maenas, by Carl Linnaeus in his 1758 tenth edition of Systema Naturae.^[30] An earlier description was published by Georg Eberhard Rumphius in his 1705 work De Amboinsche Rariteitkamer, calling the species Cancer marinus sulcatus, but this predates the starting point for zoological nomenclature.^[30] A number of later synonyms have also been published:^[30]

External identifiers for Carcinus maenas
EOL	128502
ITIS	98734
NCBI	6759
WoRMS	107381
Also found in: Wikispecies, ADW, MarLIN

• Monoculus taurus Slabber, 1778
• Cancer granarius Herbst, 1783
• Cancer viridis Herbst, 1783
• Cancer pygmaeus Fabricius, 1787
• Cancer rhomboidalis Montagu, 1804
• Cancer granulatus Nicholls, 1943
• Megalopa montagui Leach, 1817
• Portunus menoides Rafinesque-Schmaltz, 1817
• Portunus carcinoides Kinahan, 1857

The lectotype chosen for the species came from Marstrand, Sweden, but it is assumed to have been lost.^[30] In 1814, writing for The Edinburgh Encyclopaedia, William Elford Leach erected a new genus, Carcinus to hold this species alone (making it the type species of the genus, by monotypy).^[30] In 1847, Nardo described a distinct subspecies occurring in the Mediterranean Sea, which is now recognised as a distinct species, Carcinus aestuarii.^[1]

External links

Crustaceans portal

• Media related to Carcinus maenas at Wikimedia Commons
• Species Profile- European Green Crab (Carcinus maenas), National Invasive Species Information Center, United States National Agricultural Library. Lists general information and resources for European Green Crab.

References

1. ^ ^{a b} Peter K. L. Ng, Danièle Guinot & Peter J. F. Davie (2008). "Systema Brachyurorum: Part I. An annotated checklist of extant Brachyuran crabs of the world" (PDF). Raffles Bulletin of Zoology 17: 1–286. http://rmbr.nus.edu.sg/rbz/biblio/s17/s17rbz.pdf. 
2. ^ "100 of the World's Worst Invasive Alien Species". Invasive Species Specialist Group. http://www.issg.org/database/species/search.asp?st=100ss&fr=1&sts=. 
3. ^ ^{a b} S. B. Yamada & L. Hauck (2001). "Field identification of the European green crab species: Carcinus maenas and Carcinus aestuarii" (PDF). Journal of Shellfish Research 20 (3): 905–909. http://www.sgnis.org/publicat/papers/yamahauc.pdf. 
4. ^ J. V. Brian, T. Fernandes, R. J. Ladle & P. A. Todd (2005). "Patterns of morphological and genetic variability in UK populations of the shore crab, Carcinus maenas Linnaeus, 1758 (Crustacea : Decapoda : Brachyura)". Journal of Experimental Marine Biology 329 (1): 47–54. doi:10.1016/j.jembe.2005.08.002. 
5. ^ A. McKnight, L. M. Mathres, R. Avery & K. T. Lee (2000). "Distribution is correlated with color phase in green crabs, Carcinus maenas (Linnaeus, 1758) in southern New England". Crustaceana 73 (6): 763–768. doi:10.1163/156854000504787. http://www.ingentaconnect.com/content/brill/cr/2002/00000075/00000005/art00004. 
6. ^ Greg Klassen and Andrea Locke (2007). "A biological synopsis of the European green crab, Carcinus maenas". Canadian Manuscript Report of Fisheries and Aquatic Sciences No. 2818. Fisheries and Oceans Canada. http://www.dfo-mpo.gc.ca/Library/330845.pdf. 
7. ^ ^{a b} Prince William Sound Regional Citizens' Advisory Council (2004). "Non-indigenous aquatic species of concern for Alaska. Fact Sheet 1" (PDF). http://www.pwsrcac.org/docs/d0015300.pdf. Retrieved March 9, 2006. 
8. ^ E. D. Grosholz & G. M. Ruiz (1996). "Predicting the impact of introduced marine species: Lessons from the multiple invasions of the European green crab Carcinus maenas". Biological Conservation 78 (1-2): 59–66. doi:10.1016/0006-3207(94)00018-2. 
9. ^ Green Crab Control Committee (November 13, 2002). "Management plan for European Green Crab". United States Federal Aquatic Nuisance Species Task Force. http://www.anstaskforce.gov/GreenCrabManagementPlan.pdf. 
10. ^ F. J. Hidalgo et al. (2005). "A prediction come true: the green crab invades the Patagonian coast". Biological Invasions 7 (3): 547–552. doi:10.1007/s10530-004-5452-3. 
11. ^ ^{a b c} R. Thresher, C. Proctor, G. Ruiz, R. Gurney, C. MacKinnon, W. Walton, L. Rodriguez & N. Bax (2003). "Invasion dynamics of the European shore crab, Carcinus maenas, in Australia" (PDF). Marine Biology 142 (5): 867–876. http://serc.si.edu/labs/marine_invasions/publications/PDF/Thresher_et_al_2003.pdf. 
12. ^ P. J. Le Roux, G. M. Branch & M. A. P. Joska (1990). "On the distribution, diet, and possible impact of the invasive European shore crab Carcinus maenas (L.) along the South African coast". South African Journal of Marine Science 9 (1): 85–93. doi:10.2989/025776190784378835. 
13. ^ T. J. Carlton & A. N. Cohen (2003). "Episodic global dispersal in shallow water marine organisms: the case history of the European shore crabs Carcinus maenas and C. aestuarii" (PDF). Journal of Biogeography 30 (12): 1809–1820. doi:10.1111/j.1365-2699.2003.00962.x. http://www.sfei.org/bioinvasions/Reports/2003-GreenCrabDispersal383.pdf. 
14. ^ "Marine Pest Guide" (PDF). Biosecurity New Zealand. June 2006. http://www.biosecurity.govt.nz/files/pests/european-shore-crab/european-shore-crab-guide.pdf. 
15. ^ A. N. Cohen & T. J. Carlton (1995). "Introduction, dispersal and potential impacts of the green crab Carcinus maenas in San Francisco Bay, California". Marine Biology 122 (2): 225–237. http://www.springerlink.com/content/p073572hgg1185ht/. 
16. ^ J. Roman & S. R. Palumbi (2004). "A global invader at home: population structure of the green crab, Carcinus maenas, in Europe" (PDF). Molecular Ecology 13 (10): 2891–2898. doi:10.1111/j.1365-294X.2004.02255.x. PMID 15367106. http://palumbi.stanford.edu/manuscripts/Roman%20and%20Palumbi%202004.pdf. 
17. ^ ^{a b c} Perry (2005). "USGS Nonindigenous Aquatic Species Database". http://nas.er.usgs.gov/queries/FactSheet.asp?SpeciesID=190. 
18. ^ B. Bedini (2002). "Colour change and mimicry from juvenile to adult: Xantho poressa (Olivi, 1792) (Brachyura, Xanthidae) and Carcinus maenas (Linnaeus, 1758) (Brachyura, Portunidae)". Crustaceana 75 (5): 703–710. doi:10.1163/156854002760202688. 
19. ^ "European green crab". Washington Department of Fish and Wildlife. Archived from the original on June 13, 2008. http://web.archive.org/web/20080613141700/http://wdfw.wa.gov/fish/ans/identify/html/index.php?species=carcinus_maenas. Retrieved February 12, 2010. 
20. ^ M. Novak (2004). "Diurnal activity in a group of Maine decapods" (PDF). Crustaceana 77 (5): 603–620. doi:10.1163/1568540041717975. 
21. ^ E. D. Grosholz (2005). "Recent biological invasion may hasten invasional meltdown by accelerating historical introductions". PNAS 102 (4): 1088–1091. doi:10.1073/pnas.0308547102. PMC 545825. PMID 15657121. http://www.pnas.org/cgi/content/full/102/4/1088. 
22. ^ G. Miron et al. (2005). "Predation potential of the invasive green crab (Carcinus maenas) and other common predators on commercial bivalve species found on Prince Edward island" (PDF). Journal of Shellfish Research 24 (2): 579–586. http://www.sgnis.org/publicat/papers/miroaude.pdf. 
23. ^ D. L. Taylor (2005). "Predatory impact of the green crab (Carcinus maenas Linnaeus) on post-settlement winter flounder (Pseudopleuronectes americanus Walbaum) as revealed by immunological dietary analysis". Journal of Experimental Marine Biology and Ecology 324 (2): 112–126. doi:10.1016/j.jembe.2005.04.014. 
24. ^ K. D. Lafferty & K. D. Kuris (1996). "Biological control of marine pests". Ecology (Ecological Society of America) 77 (7): 1989–2000. doi:10.2307/2265695. JSTOR 2265695. 
25. ^ F. R. McEnnulty, T. E. Jones & N. J. Bax (2002). "The Web-Based Rapid Response Toolbox. Web publication". Archived from the original on May 19, 2006. http://web.archive.org/web/20060519191846/http://www.marine.csiro.au/crimp/nimpis/controlDetail.asp?ID=66. Retrieved March 9, 2006. 
26. ^ C. E. DeRivera, G. M. Ruiz, A. H. Hines & P. Jivoff (2005). "Biotic resistance to invasion: Native predator limits abundance and distribution of an introduced crab" (PDF). Ecology 86 (12): 3367–3376. doi:10.1890/05-0479. http://serc.si.edu/labs/marine_invasions/publications/PDF/DeRivera_et_al_%202005%20_Biotic_%20resistance_to_Green_Crab.pdf. 
27. ^ S. B. Yamada (2001). Global invader: the European Green Crab. Oregon Sea Grant. pp. 123. ISBN 1-881826-24-4. 
28. ^ ^{a b} J. H. R. Goddard, M. E. Torchin, A. M. Kuris & K. D. Lafferty (2005). "Host specificity of Sacculina carcini, a potential biological control agent of introduced European green crab Carcinus maenas in California" (PDF). Biological Invasions 7 (6): 895–912. doi:10.1007/s10530-003-2981-0. http://www.werc.usgs.gov/chis/Goddardetal05.pdf. 
29. ^ "FAO Nominal Catches: Carcinus maenas". FishBase. http://www.fishbase.org/report/FAO/FAOCatchList.cfm?&scientific=Carcinus%20maenas. 
30. ^ ^{a b c d e} "Species Carcinus maenas (Linnaeus, 1758), European Shore Crab". Australian Faunal Directory. Department of the Environment, Water, Heritage and the Arts. October 9, 2008. http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/Carcinus_maenas. Retrieved October 15, 2010.