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Carcinus maenas, green crab Background The brachyuran crab Carcinus maenas (Linnaeus 1758) is officially known by the common name green crab (McLaughlin et al. 2005), but other vernacular names include European green crab, shore crab and harbour crab (Williams 1984, Ingle 1983), and it is sometimes also referred to as the cockroach of the sea (http://www.oagbvg.gc.ca/domino/reports.nsf/html/c20021004se05.html). It is a member of the swimming crab family Portunidae that also includes the commercially important blue crab Callinectes sapidus covered elsewhere in this report. The green crab is native to Europe (McLaughlin et al. 2005), it’s eastern Atlantic distribution ranging from Iceland to NW Africa (Squires 1990). This species is a primary example of a rapid successful aquatic invader that has gained a strong foothold in North America. The green crab has successfully invaded both the eastern and western coasts of the USA and Canada. While the occurrence on the west coast is a more recent phenomenon, with rapid northward dispersal from San Franscico in 1989 (http://www.sei.org/crabnews.html) to British Columbia by 1998 (http://www.oagbvg.gc.ca/domino/reports.nsf/html/c20021004se05.html), the Atlantic seaboard was invaded starting at Cape Cod more than a century ago. From there the crab has since spread south to Virginia, and north into Canada, where it colonized the Bay of Fundy in the 1950’s (MacPhail 1953). It has spread along Nova Scotia in the 1980’s up to Cape Breton and into the southern Gulf of St. Lawrence by the 1990’s (Audet et al. 2003), its most northern locality in the western Atlantic. There is evidence that crabs in some localities may have originated from separate introductory events (Cameron and Metaxas 2005). In many of these areas it has established itself as the most common crab in the intertidal zone, where it is found on a variety of habitats but with a preference for protected rocky shores, cobble beaches, sandflats, and tidal marshes (Williams 1984, http://wdfw.wa.gov/fish/ans/greencrab.htm#distribution). While the life-cycle of the green crab is temperature and therefore also latitude dependent, berried females usually appear in spring (Berrill 1982). After hatching, a planktonic larval phase ensues that includes 5 stages and lasts about 2 months at 15oC (Rice and Ingle 1975), before becoming a benthic organism. Larval stages are largely believed to be the vehicle for dispersal of this species, such as by ship ballast water (http://wdfw.wa.gov/fish/ans/greencrab.htm#invasion). In New England and adjacent Canadian waters of similar temperature, the green crab matures in 2-3 years, breeds 2-3 times and has a minimum generation time of 3 years (Berrill 1982). The green crab is edible but the relatively small size (usually less than 8 cm carapace width) and yield of meat has resulted in apparently little commercial value. Thus, while it is used as bait (Elner 1985), there presently is no active fishery for this crab. However, there now are ongoing efforts for a minced-meat experimental fishery (Tremblay et al. Chapter 3 60 2006), such as in PEI (http://www.glf.dfo-mpo.gc.ca/sci-sci/crab-crab/crabwiseencrab_2002-e.html#finding_a_use) that may have the added benefit of a population control measure. Temperature limits, critical thresholds, vulnerability, and barriers to adaptation Sea surface temperatures in the current distribution of the green crab range from a February minimum of -1.8oC to an August maximum of 25.8oC. One of the reasons for the wide distribution of this crab is its tolerance to a wide range of salinities and temperatures; for the former, ranging from almost freshwater conditions of 1-3 ppt for short term exposure, and 4 ppt for longer exposure (Perkins et al. 1969,), to salinities of 54 ppt that are well above typical seawater (http://wdfw.wa.gov/fish/ans/greencrab. htm#distribution). Temperature tolerance of non-larval stages is equally impressive, ranging from close to 0°C depending on salinity, to above 30 °C, with critical thermal maxima, manifested as first signs of heat stress, ranging from 31-36 °C depending on acclimation conditions (Cuculescu et al. 1998). Not surprisingly, the overall assessment of thermal sensitivity of the green crab showed it not ranking as a sensitive species, with above average mobility and eurythermal capacity. Impact A 4oC rise in global temperature will impact the future distribution of the green crab in the western Atlantic. All models predict a loss in the southern range from Chesapeake Bay to between Delaware and Cape Cod, depending on the model. While most models predict little or no increase in range with an increase in temperature (green areas), all models indicate that the crab is already able to expand beyond its present range (crosshatching) into waters of Newfoundland and Labrador without an increase in environmental temperatures (blue areas). These areas include rocky shore habitats and prey species preferred by this species, thus not likely representing a limiting factor to such an expansion. One possible limitation is the more restricted cold tolerance of eggs and larvae compared to juveniles and adults. However, evidence is presently not clear in that regard, with successful egg incubation occurring at 10-17 oC (eggs being killed at 1.4 oC in low salinity; Wear 1974) and successful larval rearing between 10 and 25 oC (Naragaj 1993). Clearly, there is a need to understand the survival of these earlier developmental stages at temperatures between 0 and 10 oC. For example, while it is known that low salinities and absence of food at 6 oC impede or prevent larval development (http://wdfw.wa.gov/fish/ans/greencrab.htm), it is not clear how larvae would fare at that temperature under more optimal salinity and food conditions. Nevertheless, even if lower temperatures represent a present barrier for early developmental stages of the green crab, the predicted scenario of increased temperatures will increase the likelihood of a northern expansion. Thus a range expansion into more northern waters with a 4 oC temperature increase is a likely scenario. Chapter 3 61 The green crab is an omnivore (Williams 1984) capable of feeding on both animals, e.g. polychaetes and small crustaceans, and plants, i.e. seaweeds, adapting and varying its diet depending on local availability (Ropes 1968). However, it is best known as a voracious predator of molluscs, particularly bivalves of commercial importance (Elner 1981, Miron et al. 2005), such as soft-shell clams, oysters, quahogs, and mussels. The crab has negatively affected such fisheries (Congleton et al. 2005, Walton et al. 2002). Thus there is particular concern as it spreads northward and establishes large enough populations to affect clam harvests in New Brunswick and Nova Scotia, oyster production in Bras d’Or Lakes (Tremblay 2006), and the substantial blue mussel industry, as it enters the southern Gulf of St. Lawrence. Other than molluscs, there is evidence that under certain conditions the green crab can compete effectively with juveniles of commercial crustaceans, such as lobster and rock crab (Miron et al. 2005, Williams et al. 2006). There also is evidence that it is capable learning, such as improving prey-handling skills (http://wdfw.wa.gov/fish/ans/greencrab.htm#feeding). Considering these characteristics of the green crab and the abundance of commercial prey species in the areas being or likely to be invaded, an upward change in temperature will likely result in the crab affecting commercial fisheries that are presently not yet impacted as this species is “eating its way up the coast” (http://www.cbc.ca/canada/story/2002/03/16/green_crabs020316.html). References Audet, D., D.S. Davis, G. Miron, M. Moriyasu, K. Benhalima, and R. Campbell. 2003. Geographic expansion of the nonindigenous crab Carcinus maenas (L.) along the Nova Scotia shore into the southeastern Gulf of St. Lawrence, Canada. Journal of Shellfish Research 22: 255-262. Berrill, M. 1982. The life cycle of the green crab Carcinus maenas at the northern end of its range. Journal of Crustacean Biology 2: 31-39. Cameron, B., and A. Metaxas. 2005. Invasive green crab, Carcinus maenas, on the Atlantic coast and in the Bras d’Or Lakes of Nova Scotia, Canada: larval supply and recruitment. Journal of the Biological Association of the United Kingdom 85: 847855. Congleton, W.R Jr., T. Vassiliev, R.C. Bayer, and B.R. Pearce. 2005. A survey of trends in Maine soft-shell clam landings. Journal of Shellfish Research 24: 647. Cuculescu, M., Hyde, D., and K. Bowler. 1998. Thermal tolerance of two species of marine crab, Cancer pagurus and Carcinus maenas. Journal of Thermal Biology 23: 107-110. Elner, R.W. 1981. Diet of green crab Carcinus maenas (L.) from Port Hebert, Southwestern Nova Scotia. Journal of Shellfish Research 1: 89-94. Elner, R.W. 1985. Crabs of the Atlantic coast of Canada. DFO Underwater World Factsheet UW/43: 8p. Ingle, R.W. 1983. Shallow-water crabs. Cambridge University Press, London. MacPhail, J.S. 1953. Abundance and distribution of the green crab. A clam predator. Report of the Atlantic Biological Station for 1953. Fisheries Research Board of Canada, Appendix 22: 33-34. Chapter 3 62 McLaughlin, P.A, D.K. Camp, M.V. Angel, E.L. Bousfield ….. and T.T. Turgeon. 2005. Common and scientific names of auatic invertebrates from the United States and Canada: crustaceans. American Fisheries Society, Special Publication 31, Bethesda, Maryland. Miron, G., D. Audet, T. Landry and M. Moriyasu. 2005. Predation potential of the invasive green crab Carcinus maenas and other common predators on commercial bivalve species found on Prince Edward Island. Journal of Shellfish Research 24: 579-586. Nagaraj, M. 1993. Combined effects of temperature and salinity on the zoeal development of the green crab, Carcinus maenas (Linnaeus, 1758) (Decapoda: Portunidae). Scientia Marina 57: 1-8. Perkins, E.J., E. Gribbon, and R.B. Murray. 1969. Some aspects of biology of Carcinus maenas (L.). 2. Survival at low salinity. Transactions, Journal and Proceedings of the Dumfrieshire and Galloway Natural History and Antiquities Society 46: 27-28. Rice, A.L., and R.W. Ingle. 1975. The larval development of Carcinus maenas (L.) and C. mediterraneus Czerniavsky (Crustaecea, Brachyura, Portunidae) reared in the laboratory. Bulletin of the British Museum (Natural History), Zoology 28(3): 103119. Squires, H.J. 1990. Decapod Crustacea of the Atlantic Coast of Canada. Canadian Bulletin of Fisheries and Aquatic Sciences 221: 1-532. Tremblay, M.J., A. Thompson, and K. Paul. 2006. Recent trends in the abundance of the invasive green crab (Carcinus maenas) in Bras d’Or Lakes and eastern Nova Scotia based on trap surveys. Canadian Technical report of Fisheries and Aquatic Sciences 2673: iii+ 32 p. Walton W.C., C. MacKinnon, L.F. Rodriguez, C. Proctor, and G.M. Ruiz. 2002. Effect of an invasive crab upon a marine fishery: green crab, Carcinus maenas, predation upon a venerid clam, Katelysia scalarina, in Tasmania (Australia). Journal of Experimental Marine Biology and Ecology 272: 171-189. Wear, R.G. 1974. Incubation in British decapod Crustacea, and the effects of temperature on the rate and success of embryonic development. Journal of the Marine Biological Association of the United Kingdom 74: 745-762. Williams, P.J., T.A. Floyd, and M.A. Rossong 2006. Agonistic interactions between invasive green crabs, Carcinus maenas (Linnaeus), and sub-adult American lobsters, Homarus americanus (Milne Edwards). Journal of Experimental Marine Biology and Ecology 329: 66-74. Williams, A.B. 1984. Shrimps lobsters, and crabs of the Atlantic coast of the United States, Maine to Florida. Smithsonian Institution Press, Washington, D.C. Chapter 3 63