Invasion History
Panama Invasion History:
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.
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.
Taxonomy
Taxonomic Tree
Kingdom: | Animalia | |
Phylum: | Arthropoda | |
Subphylum: | Crustacea | |
Class: | Malacostraca | |
Subclass: | Eumalacostraca | |
Superorder: | Eucarida | |
Order: | Decapoda | |
Suborder: | Pleocyemata | |
Infraorder: | Brachyura | |
Superfamily: | Portunoidea | |
Family: | Portunidae | |
Genus: | Carcinus | |
Species: | maenas |
Synonyms
Cancer moenas (De Kay, 1842)
Carcinides maenas (Rathbun, 1930)
Carcinus granulatus (Smith, 1873)
Portunus maenoides (Rafinesque, 1817)
Potentially Misidentified Species
The status of the Mediterranean Green Crab, C. aestuarii Nardo 1847 (C. mediterraneus Cziernavsky 1884), as a separate species from C. maenas, has been disputed. Morphological differences between the two species include subtle differences in shapes of spines and segments, the shape of the frontal carapace, male pleopod shape, number of segments in the flagellum, and presence/absence of setae on the chelipeds. A morphometric study by Clark et al. (2001) found some overlap between the two populations. However, Yamada and Hauck (2001) listed morphological characteristics for field separation of the two species and they found an intermediate specimen from Rabat, Morocco. Genetic studies by Bagley and Geller (2000) and Roman and Palumbi (2004) support the separation of the two species.
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
OmniHabitats
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 |
Life History
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).
Regional Distribution Map
Bioregion | Region Name | Year | Invasion Status | Population Status |
---|---|---|---|---|
PAN_PAC | Panama Pacific Coast | 1866 | Non-native | Failed |
Occurrence Map
OCC_ID | Author | Year | Date | Locality | Status | Latitude | Longitude |
---|---|---|---|---|---|---|---|
3681 | Carlton and Cohen 2003 | 1866 | 1866-01-01 | None | Non-native | 8.8333 | -79.2500 |
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