Carcinus maenas

Invasion History

First Non-native North American Tidal Record: 1817
First Non-native West Coast Tidal Record: 1990
First Non-native East/Gulf Coast Tidal Record: 1817

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General Invasion History:

Carcinus maenas, also known as the Green Crab, is native to European Coasts from Iceland and Norway to Mauritania, West Africa. It has invaded six major regions, the Northwest Atlantic (from Maryland-Newfoundland), the Southwest Atlantic (Patagonia), the Southeast Atlantic (South Africa), the Northeast Pacific (from California-Alaska), the Northwest Pacific (Japan) and the Southwest Pacific (Australia) (Cohen et al. 1995; Carlton and Cohen 2003).

North American Invasion History:

Invasion History on the West Coast:

In 1989, a single Carcinus maenas was collected at Estero Americano, near Bodega Harbor, California. This may have been an isolated introduction (Grosholz and Ruiz 1995). In 1990, a specimen was caught at Redwood City, on South San Francisco Bay (Cohen et al. 1995). By 1993, C. maenas was abundant throughout the Bay (Cohen et al. 1995) and by 1994 it was found from San Francisco Bay north to Bodega Harbour (120 km) (Grosholz and Ruiz 1995). In 1997, C. maenas was collected in Coos Bay, Oregon, in 1998 it was collected in Willapa Bay, Washington, and in 1998, in Barkley Sound, British Columbia (Grozholz and Ruiz 1996; Washington Department of Fish and Wildlife 1997; Fisheries and Oceans Canada 1998). In 2007, its northern limit was Winter Harbour, Quatsino Inlet, on the west coast of Vancouver Island (50.5330 N, Klassen and Locke 2007). Populations in Barkley Sound show indications of rapid individual growth, characteristic of an expanding population (McGaw et al. 2011). In 2020, Green Crabs were collected in Haida Gwai (Queen Charlotte Islands, 53.1 Nº) and in 2021, in Ketchikan, Alaska (Miller 2022).  Recruitment in the Oregon-Washington part of the range has been sporadic, depending on favorable currents and warmer water temperatures occurring in 2003, 2005, 2006 and 2010, influenced by the Pacific Decadal Oscillation, and the El Niño Southern Oscillation (Yamada et al. 2015). Expansion into the Strait of Georgia and Puget Sound has  been slow. In 2016–2021, specimens were collected in the San Juan Islands, Padilla Bay, Dungeness Spit  and Lummi Seafood Hatchery Bay (Yamada et al. 2017, University of Washington 2017; Associated Press 2021). In May 2022, one specimen was collected in the Hood Canal;, part of Puget Sound proper (US Geological Survey Nonindigenous Species Database 2022). Establishment in Puget Sound is likely (Carolyn Tepolt, personal communication 2022). Southward dispersal has been much slower, with C. maenas spreading only to Monterey Bay and Elkhorn Slough in 1993 (Grosholz and Ruiz 1995; Ruiz et al. unpublished data) and Morro Bay in 1998, the current southern limit (USGS Nonindigenous Species Program 2009). Genetic studies indicate that the West Coast populations of C. maenas originated from the East Coast (Bagley and Geller 2000; Tepolt et al. 2009; Darling 2011). The Green Crab was likely introduced to the West Coast through the live-bait trade, or less likely, in the ballast water of ships (Cohen et al. 1995; Grosholz and Ruiz 1995). Surveys in northern California (Bodega Harbor, Bolinas Lagoon, Tomales Bay) suggest that predation by native crabs (Cancer and Metacarcinus spp.) may limit the establishment of this crab to low-salinity habitats (Jensen et al. 2007). Genetic studies suggest that the rapid spread of Green Crabs across 12 degrees of latitude in 10 years, despite low genetic diversity.  Tepolt et al. (2021) suggested that a cluster of genes in chromosomal inversion permitted local temperature adaptation despite rapid overall gene flow.

Invasion History on the East Coast:

One of the first records of Carcinus maenas from the Atlantic Coast was Say (1817), as Cancer granulosus: 'Inhabits bays and inlets near the sea'. Thomas Say is known to have collected on the Atlantic coasts of Maryland and New Jersey, so C. maenas apparently first colonized the Mid-Atlantic region, and spread north of Cape Cod in the late 19th century.

Carcinus maenas was collected by Say (1817) on the Atlantic Coast, probably in New Jersey, but possibly from the Atlantic coast of Maryland. It was also reported, as Portunus maenoides from Long Island, in 1817 (Rafinesque 1817). Other early records are from Martha’s Vineyard, Massachusetts (Gould 1841), Long Island Sound, New York, and Newport, Rhode Island (DeKay 1844). Before 1911, it was abundant at Atlantic City, New Jersey (Fowler 1911). Carcinus maenas was collected at Cape May, New Jersey by 1900 (Almaca 1963). 'Carcinus maenas, the green crab, is not common in Delaware Bay, and has only been collected by us from the Cape Henlopen tidal flat... and from the southernmost tributary entering Delaware Bay' (Leathem and Maurer 1980). Larvae were reported in plankton by Deevey 1960 (cited by Williams 1984). On the Atlantic coast of the Delmarva Peninsula, Carcinus maenas was collected in 1874 in Hog Island Bay, in Northampton Co., Virginia: 'a single male... This is the farthest south on the Atlantic coast of the United States from which this species has been reported' (Kingsley 1879). In recent surveys, this crab was found from Delaware to Virginia, in Indian River, Assawoman, Sinepuxent and Chincoteague Bays (in 2003, de Rivera et al. 2005a; Miller and Brown 2005). In 2007, a Green Crab was caught in Chesapeake Bay, in the upper Manokin River, Somerset County (Kevin Josenhans, Maryland DNR, personal communication). 

From the Gulf of Maine northwards- Carcinus maenas was collected in Provincetown, Massachusetss in 1905, and successively spread to Casco Bay, Maine in 1922-1930; Penobscot Bay (Rockland, Bar Harbor), Maine in 1951-1953; and Passamaquoddy Bay (Maine to New Brunswick Canada) and the Bay of Fundy, Nova Scotia, in 1953 (Almaca 1963; Vermeij 1982b). Contrary to some published reports, C. maenas was not collected in the Gulf of St. Lawrence until 1994, but it is now found on both the Nova Scotia and Prince Edward Island sides of Northumberland Straits, and on the north side of Prince Edward Island (Audet et al. 2003). By 1991, it crossed the Straits of Canso onto Cape Breton Island, Nova Scotia (Audet et al. 2003), and has colonized the Bras d'Or Lakes on the island (Cameron and Metaxas 2005). In 2007, C. maenas was discovered in Placentia Bay, Newfoundland (Canadian Broadcasting Company News 2007; Klassen and Locke 2007; Blakeslee et al. 2010). The expansion from Nova Scotia into the Gulf of St. Lawrence involved an introduction of genetically distinct crabs, probably of Scandinavian origin (Roman 2006; Blakeslee et al. 2010; Darling 2011). Genotypes from this second introduction are transported by currents and are appearing in populations in the Gulf of Maine (Pringle et al. 2011; Darling et al. 2014; Williams et al. 2015). The two genotypes are hybridizing in Nova Scotia and Newfoundland, but the impacts of this introgression on the crab's temperature tolerances are unknown (Jeffery et al. 2017).

In recent decades (1990s to the present), Carcinus maenas has been partially displaced from rocky shore areas, from New Jersey to Massachusetts Bay by Hemigrapsus sanguineus (Asian Shore Crab) through competition and predation on juveniles (Lohrer and Whitlatch 2002; Griffen and Delaney 2007; Griffen 2011). Carcinus maenas has been observed to move into rocky-intertidal areas at high tide, presumably from subtidal regions, or from hiding places under boulders (James Carlton, personal communication). Carcinus maenas remains common in soft-bottom habitats where H. sanguineus is absent.

Invasion History in Hawaii:

In 1873, Carcinus maenas was collected in the Hawaiian Islands (Street 1877, cited by Carlton and Cohen 2003). There are no further records from Hawaii.

Invasion History Elsewhere in the World:

In the Northwest Pacific, Carcinus spp. were recorded at Tokyo Bay in 1984. The date erroneously was given as 1958 by Sakai 1986 (cited by Carlton and Cohen 2003). Both Carcinus aestuarii (from the Mediterranean) and Carcinus maenas are present in Japan. DNA data from Bagley and Geller (2000) suggest that there was a single source population which included both species, possibly from the Iberian Peninsula, where the two species overlap (Carlton and Cohen 2003; Darling 2011). In 1999, Carcinus spp. was present in Sagami and Osaka Bays in Honshu and Dokai Bay in Kyushu, Japan (Carlton and Cohen 2003). In the Southwest Pacific, the Green Crab is established in Australia. Although regular records from New South Wales start in the 1970s, Ahyong (2005) suggests that C. maenas was established, but overlooked or misidentified, in the Sydney area since the late 19th century. In 1900, C. maenas was collected in Port Phillip Bay, Victoria (Fulton and Grant 1900), and by 1998 had spread along much of the coast of Victoria (Thresher et al. 2003). In 1993, this crab was collected on the north shore of Tasmania, and by 1998, was found in many harbors on the north and east coasts of the island. In 1976, C. maenas was collected in Adelaide Harbour, South Australia, and is established there (Furlani 1996; Thresher et al. 2003). It has not, however, become established in Western Australia's major port, Perth, where it was collected in 1965 (Furlani 1996).

In the Southern Atlantic, Carcinus spp. was collected at Table Bay Docks, Cape Town, South Africa in 1983. By 1990, Carcinus spp. ranged from Camps Bay to Cape Saldanha, a distance of 200 km (Griffiths et al. 1992). However, C. maenas failed to become established in Saldanha Bay, so that the current range is limited to Cape Town Bay and its enclosing peninsula (Robinson et al. 2005). Samples included a mix of C. maenas and C. aestuarii genotypes (Geller et al. 1997). In the Southwest Atlantic, C. maenas was collected in 2003 from Camerones Bay, Chubut Province, Patagonia, Argentina, where it is established (Hidalgo et al. 2005). Genetic studies indicate that Argentine populations were introduced from Australia (Darling 2011).

In additon to its six major established populations, Carcinus spp. (probably mostly C. maenas, but could include C. aestuarii) have been collected from many sites around the world, mostly in the tropics, where it has failed to become established, or its establishment is unknown. These sites include the Azores (Drouet 1861; Sampaio 1904, cited by Morton and Britton 2000), Rio de Janeiro and Pernambuco, Brazil (in 1857 and 1899, Carlton and Cohen 2003), the Pacific coast of Panama (in 1866, Carlton and Cohen 2003), Myanmar (Burma) (in 1933, Carlton and Cohen 2003), Ceylon (Sri Lanka) (in 1886, Carlton and Cohen 2003), Pakistan (in 1971, Carlton and Cohen 2003), the Red Sea (in 1817, Carlton and Cohen 2003), and Madagascar (in 1922, Carlton and Cohen 2003). The failure to become established in these warm waters is probably related to temperature tolerances of adult and larval stages.

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Description

The carapace of Carcinus maenas is about 3/4 long as it is broad, with a surface finely and unevenly granular, especially in the anterior half. The front has three round lobes or teeth projecting moderately between its eyes. The anterior-lateral border is slightly arched with five strong teeth, directed forward, on each side. The claws are slightly unequal, nearly smooth except for two ridges on the upper surface of the hand. The merus ('forearm') is short, while the carpus ('wrist') has a broad internal tooth or angle. The walking legs are smooth and spineless. The fifth pair of legs is slightly flattened, but is not greatly modified for swimming. The abdomen of male C. maenas is broad and triangular, with segments 3–5 fused. The mature female has a broad abdomen, with free segments. The color is highly variable, but adults are usually multicolored with a dorsal surface ranging from dark green, grayish green, or reddish, while the undersurface is yellowish white to orange. In juveniles, the color is highly variable, often with bold contrasting colors (Williams 1984).

Zoeae and megalopae larvae of C. maenas are illustrated in Roft et al. (1984) and Johnson and Allen (2005), along with additional references on larval development, and comparisons to East Coast crab larvae. Rice and Tsukimura (2007) also provide a description, with comparisons to West Coast (San Francisco Bay) crab larvae.

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Taxonomy

Ecology

General:

Life History- In crabs of the family Portunidae, the male attends the female before molting, and carries the female around, underneath his carapace. He releases the female, allows her to molt, and then copulates with her, inserting the first pair of pleopods, carrying sperm, into the female's seminal receptacles. The eggs are fertilized internally, and then extruded as a 'sponge' or a mass of eggs brooded between the abdomen and the body (Crothers 1968; Barnes 1983; Williams 1984). The number of eggs varies with size of the crab, but typical number of eggs for Carcinus maenas are around 185,000-200,000 (Broekhuysen 1936; Crothers 1966; Berrill 1982). The eggs hatch into zoea, larvae about 1 mm long, armed with long spines, which drift in the plankton. Each zoea goes through six molts, and eventually molts into a post-larval megalopa, with prominent eyes and partially developed appendages. The megalopa is capable of crawling on the bottom and active, directed swimming. After 25 to 90 days from hatching, depending on temperature and food availability, it settles and molts into a miniature 'first crab' which has all the features of an adult crab (Crothers 1968; Barnes 1983; Leignel et al. 2014).

Ecology- Carcinus maenas is most abundant in intertidal and shallow subtidal habitats through most of its range. It is more abundant in shallow, protected bays than on exposed shores (Williams 1984). In the Isles of Shoals, Gulf of Maine, Carcinus maenas is most abundant in the intertidal, while the larger Cancer irroratus (Rock Crab) and C. borealis (Jonah Crab) are most abundant in the subtidal (Donahue et al. 2009). In New South Wales, Australia, C. maenas colonized lagoons that were open to the sea > 60% of the time, and were more abundant in mangroves than in marsh or seagrass habitats (Garside et al. 2014). Fish predation may be a major factor limiting C. maenas in subtidal waters (Donahue et al. 2009).  Green crabs are omnivorous, but their diet ends to be dominated by invetebrates, espeically mollusks, crustaceans, and annelids.  Algae are usually a minor componet of the diet. (Corothers 1968;; Ropes 1989; LeRoux et al. 1990;  Rossong et al. 2011; Wong and Dowd 2014; Quinn and Boudreax 2016; Cornelius,et al. 2021; Corodone et al. 2022

Food:

molluscs; crustaceans; other inverts; algae

Consumers:

Crabs; Lobsters; Fishes, Birds

Competitors:

Trophic Status:

Omnivore

Omni

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Habitats

General Habitat	Unstructured Bottom	None
General Habitat	Grass Bed	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Salt-brackish marsh	None
General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Tidal Range	Mid Intertidal	None
Tidal Range	High Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-1	None
Maximum Temperature (ºC)	35	Experimental, Critical Temperature Maximum (CTM, rapid gradual temperatyre ncrease) varies from 32-36 C, with season and acclimation temperature (Leignel et al. 2014).Temperature Range- Equatorward distribution limited by average summer surface temperature of ~22C (Cohen et al. 1995).
Minimum Salinity (‰)	4	Salinity Range- Found in flooded tidepools in salinities as low as 1.4 ppt. 10 ppt is a more usual lower limit (Williams 1984). Larvae require at least 17-19 ppt to metamorphose and settle (Rasmussen 1973, cited by Williams 1984).
Maximum Salinity (‰)	54	Broekhuysen 1936, Experimental. Elevated salinities are possilbe in isolated tidepools, under a hot sun.
Minimum Reproductive Temperature	6	Ovigerous females, 6-10 C, Placentia Bay, Newfoundland (Best et al. 2017)
Maximum Reproductive Temperature	17	Williams 1984
Minimum Reproductive Salinity	13	Broekhuysen 1936
Maximum Reproductive Salinity	54	Broekhuysen 1936
Minimum Duration	25	Release to first crab- Berrill 1982; Lipski, unpublished data
Maximum Duration	90	Release to first crab- Berrill 1982; Lipski, unpublished data
Minimum Length (mm)	19	For adult female, 25 mm for males. Broekhuysen 1936; Crothers 1967, Crothers 1968; Berrill 1982
Maximum Length (mm)	86	For adult male, 70 mm for females, Broekhuysen 1936; Crothers 1967, Crothers 1968; Berrill 1982

General Impacts

Carcinus maenas, also known as the Green Crab, has been listed by the Invasive Species Specialist Group of the World Conservation Union (IUCN) as one of the '100 worst invasive species.'

Economic Impacts:

Fisheries- Carcinus maenas had a major impact on shellfisheries in New England, and may have similar effects on fisheries on the West Coast of North America and Australia. Predation by Green Crabs led to a reduction in Mya arenaria (Soft-Shell Clam) harvests in Maine in the 1940s-1950s (Dow and Wallace 1952). It is also a major predator on Mercenaria mercenaria (Quahog or Hard Clam) in southern New England (Walton et al. 2001). Other commercial shellfish eaten by Green Crabs in New England include blue mussels, the oyster Crassostrea virginica (Miron et al. 2005; Breen and Metaxas 2009) and Bay Scallops (Pohle et al. 1991). On the West Coast, Carcinus maenas is regarded as a potential predator on commercially important clams, including introduced Softshell Clams, Japanese Littlenecks (Venerupis philippinarum), and the Mediterranean Mussel (Mytilus galloprovincialis), as well as the native Pacific Littleneck (Leukoma staminea) (Grosholz et al. 2011). Predation by Carcinus maenas was predicted to adversely affect the fishery for the clam Katelysia scalarina in Tasmania (Walton et al. 2002). Shellfishermen use mesh bags, cultivation on ropes and in cages, to minimize predation, and may use traps to remove crabs (Walton et al. 1999; Grosholz et al. 2001). Planting large seed clams, or altering the timing of planting may also reduce losses due to Green Crab predation (Grosholz et al. 2001). Estimated current losses of bivalve fisheries (Pacific Littleneck, Japanese Littleneck, Softshell Clam, Blue Mussel) in California are negligable, but with future population increases could reach $20,000-60,000 per year (Grosholz et al. 2011).

Predation on juveniles of larger harvested crustaceans, such as American Lobster (Homarus americanus) on the East Coast (Rossong et al. 2006) and Dungeness Crab (Metacarcinus magister) on the West Coast (Cohen et al. 1995) is also a concern. However, Green Crabs are also frequent prey for large crabs and lobsters (Lynch and Rochette 2009), so impacts of the C. maenas invasion on these fisheries are difficult to determine.

In Europe, where it is native, C. maenas has long been used for food, but it is rarely caught or eaten in the United States (Williams 1984). In the US, it is widely shipped and sold as bait (Grosholz and Ruiz 1996). In Maryland and elsewhere, fishermen are encouraged to kill unused bait crabs, rather than release them, by signs posted by state agencies at fishing locations (Paul Fofonoff, personal observation).

Ecological Impacts:

Predation- On the East and West coasts of North America, and in Australia Carcinus maenas has had serious impacts on shore communities; primarily as one of the chief predators of the intertidal zone. It can affect the survival and recruitment of gastropods, bivalves, other crabs, and probably a wide range of other invertebrates (Vermeij 1982a; Vermeij 1982b; Williams 1984; Grosholz and Ruiz 2002). It is a major predator of Mya arenaria (Soft-Shell Clams) in the Gulf of Maine (Dow and Wallace 1952). In cage experiments on a mudflat in Pomquet Harbour, Nova Scotia, Carcinus maenas removed 80% of small Softshell Clams (Mya arenaria) but had negligible impacts on larger clams (Floyd and Williams 2004). In Tasmania, abundance of Carcinus maenas was negatively correlated with that of the native venerid clams Katelysia scalarina, K. rhytiphora, and Fulvia tenuicostata (Walton et al. 2002; Ross et al. 2004). On the Pacific coast, C. maenas has significantly reduced densities of the most abundant benthic taxa in Bodega Bay, California (Grosholz and Ruiz 2002).

Impacts of C. maenas's invasions are complicated by the fact that native crab species are present, and other crab species can invade, functioning as prey, competitors, and/or predators of Green Crabs. Consequently, it is necessary to compare patterns and rates of C. maenas predation to that of other crabs. In Tasmania, caging experiments showed that predation rates of C. maenas greatly exceeded those of native crabs or other predators (Walton et al. 2002). On the Oregon coast, C. maenas fed on native mussels (Mytilus trossulus) at lower rates than the native Metacarcinus magister (Dungeness Crab), but were more efficient than equal-sized Cancer magister at feeding on native Olympia Oysters, Ostrea lurida, because of greater claw strength (Yamada and Kosro 2010). In the Bras d'Or Lakes, Nova Scotia, Breen and Metaxas (2009) measured predation rates of juvenile and adult C. maenas on mussels (Mytilus sp.) and compared them to two species of native crabs Cancer irroratus (Rock Crab) and Dyspanopeus sayi (Say's Mud Crab). Rates of mussel consumption by C. maenas were similar or lower than those of the native species, but a favorable year for recruitment could increase the crabs' impact (Breen and Metaxas 2009). In field experiments at Avery Point, Long Island Sound, C. maenas fed on young mussels at a higher rate than the recently introduced crab, Hemigrapsus sanguineus (Asian Shore Crab), but the much higher densities of H. sanguineus now make it the more important predator in the rocky intertidal south of Cape Cod (Lohrer and Whitlach 2002).

Interactions among crabs of different sizes often result in predation, either among crabs of the same species or different species. Over an 11-year period in Bodega Harbor, Hemigrapsus oregonensis abundance was negatively correlated with C. maenas abundance, but recovered, with a lag period, when C. maenas declined (de Rivera et al. 2011). Large C. maenas prey on small H. sanguineus and vice versa (Griffen and Byers 2009). Predation, aggression, and interference behavior have the effect of reducing the predation rates of both species when they co-occur. In laboratory experiments, Carcinus maenas was found to consume juvenile lobsters in 6 of 11 trials (Rossong et al. 2006). Further, very small lobsters (under 35 mm carapace length) showed reduced foraging in the presence of Green Crabs. However, field studies and laboratory experiments indicate that rates of predation are low, and that predation on C. maenas by lobsters may be equally or more frequent (Lynch and Rochette 2009).

The invasion of a new predator, such as C. maenas, can also alter the behavior and morphology of prey species. In a system of tidepools at Nahant, Littorina littorea (the common Periwinkle) responded to increased C. maenas density by moving to other pools (Trussell et al. 2004). Softshell Clams (Mya arenaria) in the Damariscotta and Wells estuaries, Maine, burrowed deeper in the bottom sediment in the presence of C. maenas, responding both to chemical and mechanical signals (Whitlow et al. 2003; Flynn and Smee 2010). Whitlow (2010) found that chemical cues induced both deeper burrowing and growth of longer siphons in the clams. In Bodega Harbor, Calfiornia, predation by C. maenas resulted in reduced use of the lower intertidal zone by the native crab Hemigrapsus oregonensis, an effect that persisted, even after the abundance of C. maenas declined (de Rivera et al. 2011).

The invasion of Green Crabs has resulted in evolutionary changes in some prey populations, and in a possible ‘arms race’ as C. maenas has responded to these changes in its prey. For several species of gastropods in the Gulf of Maine, selective predation by C. maenas has apparently resulted in changes in shell morphology which make the shells more resistant to crushing. This has been shown for Littorina obtusata (Seeley 1986; Edgell et al. 2009; Edgell and Hollander 2011) and Nucella lapillus (Vermij 1982a). The picture for N. lapillus has been complicated by an overall increase in shell size over the last 80 years, which accounts for the increase in thickness when corrected for allometry, which could be a response to predation, or due to other causes (Fisher et al. 2009). This increase in shell strength, greatest in more southern populations, which have coexisted with C. maenas for a longer time, has been partially compensated for by an increase in crusher claw size and strength in more southern populations of C. maenas (Smith 2004; Edgell and Rochette 2008). The morphological response of Carcinus' crusher claw may be limited in northern populations by metabolic effects of temperature (Baldridge and Smith 2008).

These temporal and geographical differences have not been found in Littorina littorea, possibly because of the long-range dispersal of its planktonic larvae (Vermeij 1982b) or because of its ancestral co-occurrence with C. maenas (Edgell and Rochette 2008). Edgell and Rochette (2008) found that Carcinus claw scars and shell damage were less frequent on L. littorea than on L. obtusata.

Competition- Carcinus maenas is a potential competitor with native crabs, but this has not been well-studied on the Atlantic coast. In experiments on antagonistic behavior, the largest crab usually wins, which favors Callinectes sapidus (Blue Crab) because of its larger adult size (Ruiz et al. unpublished data; de Rivera et al. 2005). In competition for food, C. maenas may have a disadvantage against the faster-moving swimming crabs (Callinectes sapidus, Ovalipes ocellatus- Calico Crab) (Ropes 1989; Ruiz et al. unpublished data). Competition may restrict the penetration of C. maenas into estuarine habitats favored by C. sapidus. Interference competition and aggression occur between Carcinus maenas and Hemigrapsus sanguineus. In experiments, interference between the two species lowered the predation rates of both species on amphipods (Griffen and Byers 2006). In Tasmania, the invasion of Carcinus maenas apparently resulted in the displacement of the native crab Pachygrapsus gaimardi (Ruiz et al. unpublished). Caging experiments in King Georges Sound, Tasmania, showed apparent competition between C. maenas and the introduced starfish Asterias amurensis. The two species overlapped in depth range, and prey choice, but showed partitioning, with C. maenas preferring shallower water and smaller clams.

Food/Prey- Breen and Metaxas (2009) found little evidence of competition when juvenile C. maenas, and native Rock Crabs (Cancer irroratus) of similar size were reared together. Instead, the growth rate of C. irroratus increased, as a result of feeding on green crabs (Breen and Metaxas 2009).

Trophic Cascades- As a novel top predator in many littoral ecosystems, C. maenas invasions have resulted in effects across several trophic levels, affecting organisms which do not interact directly with the crabs. In a system of tidepools at Nahant, Massachusetts Bay, Littorina littorea (Common Periwinkle) responded to increased C. maenas density by moving to other pools. The reduction of grazing resulted in increased growth of ephemeral red and green algae (Trussell et al. 2004). Similarly, in the Gulf of Maine, Carcinus maenas preyed intensely on the sea slug Placida dendritica, which grazes on the introduced alga Codium fragile, which could favor the growth of the seaweed in Green Crab habitats, such as enclosed harbors and estuaries (Harris and Jones 2005). After the Carcinus maenas invasion in Bodega Bay Harbor, California, several invertebrate species, including the polychaetes Exogene sp. and Lumbrinereis sp. and the tanaid Leptochelia dubia increased in abundance, probably as an indirect effect of reduction in Nutricola spp. populations (Grosholz et al. 2000). The introduced clam Gemma gemma increased dramatically (two orders of magnitude) after the Carcinus invasion, apparently because of decreased competition from native Nutricola clams (Grosholz 2005). In nearby Tomales Bay, the invasion of C. maenas combined with that of the introduced Atlantic Oyster Drill (Urosalpinx cinerea) nearly eliminated the native Olympic Oyster (Ostrea lurida) from the inner, low-salinity, region of Tomales Bay (Kimbro et al. 2009).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Predation
In Bodega Harbor CA, abundances of the clams Nutricola tantilla and N. confusa and the native crab Hemigrapsus oregonensis were sharply reduced after the invasion of Carcinus maenas. Experiments indicated high rates of feeding on the native clams and crabs (Grosholz et al. 2000). Over an 11-year period in Bodega Harbor, H. oregonensis abundance was negatively correlated with C. maenas abundance, but recovered, with a lag period, when C. maenas declined. However, C. maenas predation had persisting effects on the size and intertidal distribution of H. oregonensis (de Rivera et al. 2011). In Tomales Bay, Carcinus maenas is a less effective predator than the native crab (Cancer antennarius), on native (Acanthinucella spirata) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Competition
In laboratory experiments, Carcinus maenas spent more time around bivalve bait than the native crab Hemigrapsus oregonensis, and was more successful at approaching competitors surrounding the bait than H. oregonensis, in Bodega Bay Harbor CA (Jensen et al. 2002). Carcinus maenas also displaces Metacarcinus magister (Dungeness Crabs) of equal size in feeding trials, and causes M. magister to emigrate from shelters in laboratory trials (McDonald et al. 2001). In enclosure experiments, Green Crabs reduced the abundance and quality of food available for wintering shorebirds (Dunlin, Calidris alpina) (Estelle and Grosholz 2012).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Trophic Cascade
After the Carcinus maenas invasion in Bodega Bay Harbor, California, several invertebrate species, the polychaetes Exogene sp. and Lumbrinereis sp. and the tanaid Leptochelia dubia increased in abundance, probably as an indirect effect of reduction in Nutricola spp. populations (Grosholz et al. 2000). The introduced clam Gemma gemma increased dramatically (two orders of magnitude) after the Carcinus invasion, apparently because of decreased competition from native Nutricola clams (Grosholz 2005). In Tomales Bay, Carcinus maenas is a less effective predator than the native crab, Cancer antennarius, on native (Acanthinucella spirata) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009). The increased abundance of U. cinerea, also more tolerant of low salinity than native whelks, has resulted in increased mortality and a near-absence of the native Olympia Oyster (Ostrea lurida).
P110	Tomales Bay		Ecological Impact	Predation
In Tomales Bay, Carcinus maenas is a less effective predator than the native crab, Cancer antennarius (California Rock Crab), on native (Acanthinucella spirata) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009).
P110	Tomales Bay		Ecological Impact	Trophic Cascade
In Tomales Bay, Carcinus maenas is a less effective predator than the native crab, Cancer antennarius, on native (Acanthinucella spirata, Angular Unicorn Whelk) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009). The increased abundance of U. cinerea, also more tolerant of low salinity than native whelks has resulted in increased mortality and a near-absence of the native Olympia Oyster (Ostrea lurida).
P112	_CDA_P112 (Bodega Bay)		Ecological Impact	Predation
In Bodega Harbor CA, abundances of the clams Nutricola tantilla and N. confusa and the native crab Hemigrapsus oregonensis were sharply reduced after the invasion of Carcinus maenas. Experiments indicated high rates of feeding on the native clams and crabs (Grosholz et al. 2000). Over an 11-year period in Bodega Harbor, H. oregonensis abundance was negatively correlated with C. maenas abundance, but recovered, with a lag period, when C. maenas declined. However, C. maenas predation had persisting effects on the size and intertidal distribution of H. oregonensis (de Rivera et al. 2011).
P112	_CDA_P112 (Bodega Bay)		Ecological Impact	Competition
In laboratory experiments, Carcinus maenas spent more time at bivalve baits than the native crab Hemigrapsus oregonensis, and was more successful at approaching competitors surrounding bait than H. oregonensis, in Bodega Harbor California (Jensen et al. 2002). Carcinus maenas also displaces Metacarcinus magister (Dungeness Crabs) of equal size in feeding trials, and causes M. magister to emigrate from shelters in laboratory trials (McDonald et al. 2001). In enclosure experiments, Green Crabs reduced the abundance and quality of food available for wintering shorebirds (Dunlin, Calidris alpina) (Estelle and Grosholz 2012).
P112	_CDA_P112 (Bodega Bay)		Ecological Impact	Trophic Cascade
After the Carcinus maenas invasion in Bodega Bay Harbor, California, several invertebrate species, specifically the polychaetes Exogene sp. and Lumbrinereis sp. and the tanaid Leptochelia dubia increased in abundance, probably as an indirect effect of reduction in Nutricola spp. populations (Grosholz et al. 2000). The introduced clam Gemma gemma increased dramatically (two orders of magnitude) after the Carcinus invasion, apparently because of decreased competition from native Nutricola clams (Grosholz 2005).
NEP-V	Northern California to Mid Channel Islands		Economic Impact	Fisheries
In Bodega Bay, Caliornia, Manila Clams (Venerupis philippinarum) planted in mesh bags, were prone to heavy predation by Carcinus maenas. Predation was reduced by planting the clams later in the season, when the clams were larger (Grosholz et al. 2001). Estimated current losses of bivalve fisheries (Pacific Littleneck, Japanese Littleneck, Softshell Clam, Blue Mussel) in California are negligable, but with future population increases of C. maenas, could reach $20,000-60,000 per year (Grosholz et al. 2011).
P112	_CDA_P112 (Bodega Bay)		Economic Impact	Fisheries
In Bodega Bay, California, Manila Clams (Venerupis philippinarum) planted in mesh bags, were prone to heavy predation by Carcinus maenas. Predation was reduced by planting the clams later in the season, when the clams were larger (Grosholz et al. 2000).
NEP-IV	Puget Sound to Northern California		Ecological Impact	Predation
Carcinus maenas fed at lower rates overall than native Metacarcinus magister (Dungeness Crab), collected from the Oregon coast (no location specified), when feeding on native mussels (Mytilus trossulus), but were more efficient than equal-sized M. magister at feeding on native Olympia Oysters, Ostrea lurida, because of greater claw strength (Yamada et al. 2010). Palacios and Ferraro (2003) found that Carcinus maenas preferred O. lurida to 3 other species of bivalves (introduced Venerupis philippinarum (Japanese Littleneck) and native Macoma nasuta (Bent-nose Macoma) and Cryptomya californica (California Softshell), when offered in equal amounts.
CA	California		Ecological Impact	Competition
In laboratory experiments, Carcinus maenas spent more time around bivalve bait than the native crab Hemigrapsus oregonensis, and was more successful at approaching competitors surrounding the bait than H. oregonensis, in Bodega Bay Harbor CA (Jensen et al. 2002). Carcinus maenas also displaces Metacarcinus magister (Dungeness Crabs) of equal size in feeding trials, and causes M. magister to emigrate from shelters in laboratory trials (McDonald et al. 2001). In enclosure experiments, Green Crabs reduced the abundance and quality of food available for wintering shorebirds (Dunlin, Calidris alpina) (Estelle and Grosholz 2012).  In laboratory experiments, Carcinus maenas spent more time at bivalve baits than the native crab Hemigrapsus oregonensis, and was more successful at approaching competitors surrounding bait than H. oregonensis, in Bodega Harbor California (Jensen et al. 2002). Carcinus maenas also displaces Metacarcinus magister (Dungeness Crabs) of equal size in feeding trials, and causes M. magister to emigrate from shelters in laboratory trials (McDonald et al. 2001). In enclosure experiments, Green Crabs reduced the abundance and quality of food available for wintering shorebirds (Dunlin, Calidris alpina) (Estelle and Grosholz 2012).
CA	California		Ecological Impact	Predation
In Bodega Harbor CA, abundances of the clams Nutricola tantilla and N. confusa and the native crab Hemigrapsus oregonensis were sharply reduced after the invasion of Carcinus maenas. Experiments indicated high rates of feeding on the native clams and crabs (Grosholz et al. 2000). Over an 11-year period in Bodega Harbor, H. oregonensis abundance was negatively correlated with C. maenas abundance, but recovered, with a lag period, when C. maenas declined. However, C. maenas predation had persisting effects on the size and intertidal distribution of H. oregonensis (de Rivera et al. 2011). In Tomales Bay, Carcinus maenas is a less effective predator than the native crab (Cancer antennarius), on native (Acanthinucella spirata) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009)., In Tomales Bay, Carcinus maenas is a less effective predator than the native crab, Cancer antennarius (California Rock Crab), on native (Acanthinucella spirata) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009)., In Bodega Harbor CA, abundances of the clams Nutricola tantilla and N. confusa and the native crab Hemigrapsus oregonensis were sharply reduced after the invasion of Carcinus maenas. Experiments indicated high rates of feeding on the native clams and crabs (Grosholz et al. 2000). Over an 11-year period in Bodega Harbor, H. oregonensis abundance was negatively correlated with C. maenas abundance, but recovered, with a lag period, when C. maenas declined. However, C. maenas predation had persisting effects on the size and intertidal distribution of H. oregonensis (de Rivera et al. 2011).
CA	California		Ecological Impact	Trophic Cascade
After the Carcinus maenas invasion in Bodega Bay Harbor, California, several invertebrate species, the polychaetes Exogene sp. and Lumbrinereis sp. and the tanaid Leptochelia dubia increased in abundance, probably as an indirect effect of reduction in Nutricola spp. populations (Grosholz et al. 2000). The introduced clam Gemma gemma increased dramatically (two orders of magnitude) after the Carcinus invasion, apparently because of decreased competition from native Nutricola clams (Grosholz 2005). In Tomales Bay, Carcinus maenas is a less effective predator than the native crab, Cancer antennarius, on native (Acanthinucella spirata) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009). The increased abundance of U. cinerea, also more tolerant of low salinity than native whelks, has resulted in increased mortality and a near-absence of the native Olympia Oyster (Ostrea lurida)., In Tomales Bay, Carcinus maenas is a less effective predator than the native crab, Cancer antennarius, on native (Acanthinucella spirata, Angular Unicorn Whelk) and introduced whelks (Urosalpinx cinerea, Atlantic Oyster Drill), resulting in an increased abundance and habitat range of whelks in the inner Bay, where C. maenas better tolerates low salinities (Kimbro et al. 2009). The increased abundance of U. cinerea, also more tolerant of low salinity than native whelks has resulted in increased mortality and a near-absence of the native Olympia Oyster (Ostrea lurida)., After the Carcinus maenas invasion in Bodega Bay Harbor, California, several invertebrate species, specifically the polychaetes Exogene sp. and Lumbrinereis sp. and the tanaid Leptochelia dubia increased in abundance, probably as an indirect effect of reduction in Nutricola spp. populations (Grosholz et al. 2000). The introduced clam Gemma gemma increased dramatically (two orders of magnitude) after the Carcinus invasion, apparently because of decreased competition from native Nutricola clams (Grosholz 2005).
CA	California		Economic Impact	Fisheries
In Bodega Bay, Caliornia, Manila Clams (Venerupis philippinarum) planted in mesh bags, were prone to heavy predation by Carcinus maenas. Predation was reduced by planting the clams later in the season, when the clams were larger (Grosholz et al. 2001). Estimated current losses of bivalve fisheries (Pacific Littleneck, Japanese Littleneck, Softshell Clam, Blue Mussel) in California are negligable, but with future population increases of C. maenas, could reach $20,000-60,000 per year (Grosholz et al. 2011)., In Bodega Bay, California, Manila Clams (Venerupis philippinarum) planted in mesh bags, were prone to heavy predation by Carcinus maenas. Predation was reduced by planting the clams later in the season, when the clams were larger (Grosholz et al. 2000).

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