Invasion History

First Non-native North American Tidal Record: 1840
First Non-native West Coast Tidal Record: 1968
First Non-native East/Gulf Coast Tidal Record: 1840

General Invasion History:

Littorina littorea is native to Europe from the White Sea, Russia to Gibraltar, but absent (except as failed introductions) from the Mediterranean (Bequaert 1943; Reid 1998). The first collection of living animals on the Atlantic Coast of North America was in 1840 at Pictou, Nova Scotia (Willis 1857, cited by Ganong 1887). Although its status has been disputed (Clarke and Erskine 1963; Wares et al. 2002), general historical records and recent molecular data strongly support its introduction to the Atlantic coast of North America sometime in the last 1,000 years, possibly with multiple introductions (Spjeldnaes and Henningsmoen 1963; Blakeslee et al. 2008; Chapman et al. 2008; Brawley et al. 2009). On the west coast of North America, scattered specimens are known from Newport Bay, California to Puget Sound, Washington but establishment of populations has not been documented (Hanna 1966; Carlton 1969; Carlton 1979; Carlton 1992; Chang et al. 2011).

North American Invasion History:

Invasion History on the West Coast:

Scattered occurrences of Littorina littorea have been noticed from California to Washington since 1937. Early occurrences in Puget Sound, Washington in 1937 (Hanna 1966), and Trinidad Bay, California (Carlton 1969) may have resulted from transplants of Eastern Oysters (Crassostrea virginica). Littorina littorea was found in San Francisco Bay in 1968-70 and 1977, but did not become established (Carlton 1992). Another specimen was found in 1995 (Cohen and Carlton 1995), and occasional occurrences have been noted since (Ruiz et al. unpublished data; Chang et al. 2011). In southern California, a single shell was collected in Newport Bay in 1975 (Carlton 1979). A small patch of abundant snails was found in Anaheim Bay in 2002 and was eradicated, apparently successfully (Chang et al. 2011). Several populations of L. littorea in San Francisco Bay were examined, and found to consist entirely of adult snails, with high diversity of genotypes typical of the East Coast, strongly suggesting that these animals had failed to reproduce and had not undergone the 'genetic bottleneck' of reduced diversity typical of newly established invasive populations (Chang et al. 2011). In 2010, populations of L. littorea were found at two sites in Burrard Inlet, Vancouver, British Columbia (354 animals, all adults). These were similar in size to snails purchased from local markets, but had been resident on the shore for some time and had epibionts including algae and barnacles. The periwinkles apparently have not become established, as none were collected after August 2011 (Harley et al. 2013).

Invasion History on the East Coast:

Living specimens of Littorina littorea were first collected on the East Coast at Pictou, Nova Scotia (Willis 1857, cited by Ganong 1887), but they rapidly became one of the most abundant littoral gastropods on Northeastern shores (Verrill 1880; Ganong 1887). As noted above, there has been some uncertainty about the native/introduced status of L. littorea on the East coast of North America. A single fossil specimen from ~40,000 B.P. is known from Nova Scotia (Wagner 1977, cited by Reid 1996). Specimens of L. littorea (dating back to ~1000-1300 B.C.) have been found in Nova Scotia and Newfoundland in Indian and Norse archaeological sites (Clarke 1961; Clarke and Erskine 1963; Spjeldnaes and Henningsmoen 1963). This has been taken to infer the presence of native populations of L. littorea in eastern Canada (Clarke and Erskine 1963) or the introduction of this species by Viking explorers (Spjeldnaes and Henningsmoen 1963). One recent molecular analysis (nuclear and mitochondrial DNA) suggested that populations of Littorina littorea on the Atlantic coast of North America are native (Wares et al. 2002). It has been argued that oceanographic conditions prevented the spread of L. littorea until their rediscovery around 1840 (Clarke and Erskine 1963; Wares et al. 2002). However, the combination of historical, paleontological, and genetic evidence supports introduced status for L. littorea (Chapman et al. 2007).

A more comprehensive genetic analysis by Blakeslee et al. (2008), incorporating molecular analysis with a greater number of sampled haplotypes, parasitological studies, and more sophisticated statistical analysis strongly supports an introduced status for L. littorea in North America and for its common trematode parasite, Cryptocotyle lingua. Other evidence for its introduced status includes: the absence of L. littorea shells in natural deposits, the lack of early collections of a relatively large (and edible) snail, and the improbability of a sudden range expansion, from a small range in Atlantic Canada to ~1,000 km of coastline, in about 50 years (Chapman et al. 2007). The genetic diversity of L. littorea suggests multiple introductions, possibly as early as Viking times, based on estimates using genetic divergence and assumed mutation rates (400 to 1,000 years ago), but with wide confidence intervals. Brawley et al. (2009) examined more closely historical shipping records at Pictou, Nova Scotia, a major port for immigrants from Great Britain and Ireland, and compared genotypes of Pictou and British Isles populations. They estimated divergence times with a lower confidence limit of 192-384 years, roughly comparable with a late 18th-early 19th century introduction suggested by shipping records (Brawley et al. 2009).

From the first collection at Pictou, Nova Scotia in 1840, on the southern Gulf of St. Lawrence, the known range of L. littorea expanded rapidly to the north and south. In 1855, it was collected in Bathurst. New Brunswick, on the Bay of Chaleur (Morse 1880, cited by Bequaert 1943), and in 1882 in Lake Melville, Labrador (USNM 34326, US National Museum of Natural History, 54°N; 60°W, collected by Stearns). This appears to be the current northern limit for this species (Bequaert 1943; Reid 1996). In 1857, it was collected in Halifax, on the Atlantic Coast of Nova Scotia, and in the Bay of Fundy in 1861 (Ganong 1887). By 1873, it occurred at Saco, Maine 'in abundance', and reached Provincetown, Massachusetts by 1870, at which time it was 'very rare'. By 1875, it was abundant there (Bequaert 1943; Dexter; 1961).

To the south of Cape Cod, in 1875 L. littorea was rare at Woods Hole, Massachusetts, but abundant by 1876 (Ganong 1887). Its spread to the south was rapid: New Haven, Connecticut in 1879 (Bequaert 1943); Newport, Rhode Island in 1880 (Ganong 1887); Staten Island, New York in 1888 (Bequaert 1943); Atlantic City, New Jersey in 1892 (Bequaert 1943). However, the range expansion slowed down sharply, south of Atlantic City. Littorina littorea reached Cape May, New Jersey by 1928 (Bequaert 1943) and Cape Henlopen, Delaware by 1970 (Kraueter 1974). In 1959, Wells (1965) collected L. littorea in the Maryland region, stating it was rare 'attached to the rocks of a jetty at West Ocean City, directly opposite Ocean City Inlet'. At Assateague Island, Maryland-Virginia in 1988-1989 (exact site not given), it was found 'only on rock jetties and wooden groins' (Counts and Bashore 1991). It was reported from Chincoteague, Virginia in 1971 (Kraueter 1974) and later at Wachapreague (date not given) (Vermeij 1982b).

Invasion History Elsewhere in the World:

In the Mediterranean, Littorina littorea has been collected in Italy, in the Lagoon of Faro, Sicily in 1978 and in Livorno, on the Tyrrhenian Sea, but these populations were extinct by 1988 (Johanesson 1988; Zenetos et al. 2004; Crocetta 2012).


Description

The shell of Littorina littorea is solid, heavy and turban shaped, topped with an onion-shaped spire. It is dextrally coiled, with 6 or 7 whorls, and lacks an umbilicus. The shell surface has shallow sutures, resulting in flat-sided whorls on the spire, sculpted by low, narrow ribs and fine spiral lines. The color varies from uniform cream to brown, black, orange, or red, but usually pale brown with 8-25 narrow black or brown lines. The columella is white and the outer edge of the aperture is brown or with brown lines. The interior of the aperture is brown. The shell commonly reaches about 31 mm in length, but the maximum is 53 mm. Description from: Bequaert 1943; Morris 1975; Gosner 1978; and Reid 1996.

This species lays its eggs in distinctive transparent, planktonic, egg capsules, in the form of flat disks (0.3 - 1.1 mm diameter) (Reid 1998). The planktotrophic veligers are distinguished by a prominent red or black pigmented spot on the anterior corner of each lobe of the velum. The veligers reach about 0.6 mm before settling (Newell and Newell 1997).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Gastropoda
Order:   Neotaenioglossa
Family:   Littorinidae
Genus:   Littorina
Species:   littorea

Synonyms

Potentially Misidentified Species


None

Littoraria irrorata
None

Littorina saxatilis
None

Ecology

General:

Littorina littorea occurs on rocky coasts, but also on mud, sand and marsh habitats in some parts of its range. Sexes are separate, with males slightly larger than females, and fertilization is internal. This snail reaches a mature size in one year and has a life span of at least 5 years. Aquarium specimens are reported to have lived 10 years or more (Clay 1961). The eggs are laid in water and are contained in gelatinous capsules, in the shape of a shallow dome, surrounded by a flat brim, each containing 1-9 (but usually 1-3) eggs. The spawning season varies with latitude, from summer in the Barents Sea to late winter-spring or early summer in Denmark, England, and the Netherlands (Clay 1961; Reid 1996). In New Brunswick, spawning occurs in April to November (Chase and Thomas 1995a), while in Narragansett Bay, Rhode Island egg capsules and larvae occurred year-round, but were rare in July-August (Fofonoff, unpublished data). Females in New Brunswick released an estimated 50,000 to 200,000 embryos per season. The egg capsules are usually suspended in the plankton and can be abundant in nearshore waters. The eggs take about 6 days to hatch into planktotrophic veligers, which remain in the plankton for 2-4 weeks (Fretter and Graham 1962). The larvae settle with shells about 0.275 mm in length and 0.375-390 mm in diameter (Reid 1996).

Juveniles and adults of the Common Periwinkle inhabit the middle and upper intertidal zone, primarily of rocky shores, but also extending into mudflats and marshes. On rocky shores, their distribution is often associated with rockweeds (Fucaceae) and is affected by wave action and food availability (Clay 1961; Reid 1996). The preference of L. littorea for the mid- and upper intertidal zone is related to the presence of more effective predators in the subtidal zone. Growth of caged juvenile snails in the Gulf of Maine, was greater in the subtidal and lower intertidal, but a subtidal predator, the Jonah Crab (Cancer borealis), was a more effective predator on larger snails than the intertidal Green Crab (Carcinus maenas) (Perez et al. 2009). However, some specimens of L. littorea have been found at maximum depths ranging from 15 m to 60 m, especially in more northern locations (Reid 1996). On soft sediments, in sheltered areas, periwinkles are usually associated with scattered stones, shells, mussel beds, or wood, which provide sites of attachment. The snails leave these refuges to make feeding excursions on the sediment surface (Reid 1996).

As an inhabitant of the temperate intertidal zone, the Common Periwinkle is tolerant of a wide range of temperature and salinity. It can tolerate temperatures as high as 41°C for up to 48 h (Fraenkel 1960), but an average summer temperature of 21°C seems to mark the southern limit for this species (Wells 1965). It can also tolerate short-term exposure to fresh water from rain and runoff, but reported lower limits for long-term survival range from 9.5 to 16 PSU (Todd 1964; Remane and Schleiper 1971; Reid 1996). Todd (1964) found a strong effect of temperature on salinity tolerance, with animals surviving well at 11 PSU at 5°C, but dying at salinities below 16 PSU at 15°C. The lower salinity limit for reproduction was about 15 PSU, in Kiel Bay, Germany, but was around 20 PSU in other European locations (Hayes 1929, cited by Remane and Schlieper 1971).

Littorina littorea is an omnivorous grazer, feeding on a wide range of algae and on small attached organisms, especially opportunistic algae, such as Ulva, Cladophora and Ectocarpus, and early settlement stages (germlings) of more robust seaweeds such as Fucus (Lubchenco 1978; Lubchenco and Menge 1978; Lubchenco 1983; Bertness et al. 1984; Reid 1996; Bertness et al. 2004). On soft sediment, L. littorea grazes benthic microalgae on the sediment surface and also grazes on the leaves and rhizomes of Spartina alterniflora (Smooth Cordgrass), the dominant marsh grass in northeastern salt marshes (Reid 1996; Bertness 1984; Tyrell et al. 2008). This periwinkle is basically a herbivore, but does ingest small animals, such as newly attached barnacles and mussels, and the egg capsules of other snails, such as those of Ilyanassa obsoleta. Whether it digests these organisms is unknown (Brenchley 1982; Brenchley and Carlton 1983; Reid 1996).

As a very abundant intertidal organism on the coast of Northeastern North America, L. littorea is an important component of coastal food webs. It is a major food item for crabs, fishes, and shorebirds, and also a major final and intermediate host of numerous parasites. Among frequent predators in East Coast waters are Green Crabs (Carcinus maenas), Asian Shore Crabs (Hemigrapsus sanguineus), Blue Crabs (Callinectes sapidus), Jonah Crabs (Cancer borealis), Dogwinkles (Nucella lapillis), Oyster Drills (Urosalpinx cinereus), Mummichogs (Fundulus heteroclitus), sandpipers (Calidris spp.), gulls (Larus spp.), and crows (Corvus spp.) (Dexter 1947; Peterson 1979; Gerard et al. 1999; Trussell et al. 2004; Ellis et al. 2007; Byers et al. 2008; Perez et al. 2009).

The Common Periwinkle hosts a variety of parasites. Eleven species of digenean trematodes are known from Europe, while only six have been found in Eastern North American populations (Blakeslee and Byers 2008). The most abundant of these, Cryptocotyle lingua, was introduced to North American waters, probably with L. littorea, based on genetic data (Blakeslee et al. 2008). The rediae (first parasitic stage) of C. lingua occur at an average prevalence of 9-10% in North American L. littorea populations, but can be much more abundant locally. Prevalence at New England intertidal sites was strongly influenced by the abundance of Herring Gulls (Larus argentatus) the final host of the parasite (Byers et al. 2008). Heavy infection with these parasites can damage the snail's digestive organs and greatly reduce their grazing and reproductive rates (Blakeslee and Byers 2008). In addition to obvious parasites, the bodies and shells of introduced L. littorina (sampled in Nova Scotia) host a wide variety of endosymbionts (at least 7 phyla), whose exact identity, native status, and impacts on the snail's biology are unknown (Buckland-Nicks et al. 2013).

Food:

benthic algae; detritus; egg capsules

Consumers:

crabs, fishes, shorebirds, humans

Competitors:

Trophic Status:

Herbivore

Herb

Habitats

General HabitatUnstructured BottomNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatSalt-brackish marshNone
Tidal RangeLow IntertidalNone
Tidal RangeMid IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)0.1Field Data- Denmark (Clay 1961)
Maximum Temperature (ºC)41Survival - Lethal temperature from 48 hour experiment. (Fraenkel 1960). There is a relationship between lethal temperature and the level of shore inhabited by snail- Periwinkles from high tide levels have higher lethal temperatures than those from low tide level; also varies seasonally (Clay 1961). Although this snail tolerates very high short-term temperatures, such as might be found in tide pools, an average summer temperature of 21 C seems to mark the southern limit for this species (Wells 1965).
Minimum Salinity (‰)9.5Field: Remane and Schleiper 1971; Reid 1996. Todd (1964) found that survival at low salinities varied with temperature, with lower limits at 11 PSU at 5 C, but 16 at 15 C.
Maximum Salinity (‰)48Experimental (Todd 1964)
Minimum Reproductive Salinity15Field, Kiel Bay, Germany. However, 20 PSU is a more common limit for populations outside the Baltic (Hayes 1929, cited by Remane and Schleiper 1971)
Minimum Duration20Planktonic egg (6 days, typical) + Larval Period (Fretter and Graham 1962)
Maximum Duration34Planktonic egg (6 days, , typical) + Larval Period (Fretter and Graham 1962)
Minimum Length (mm)10.6Minimum size at maturation (Reid 1996)
Maximum Length (mm)53Maximum reported size, but a more usual maximum is ~31 mm (Bequaert 1943; Morris 1975; Gosner 1978; Reid 1996)
Broad Temperature RangeNoneCold-Temperate
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Littorina littorea is a common food item in Europe, but is rarely eaten in the US. However, it has had major impacts on food webs, parasite communities, and even on the physical structure of coastlines in Northeastern North America. It is often one of the most abundant grazers in rocky intertidal areas, where it can control the abundance and composition of seaweeds. It is also abundant in marshes, mudflats, and cobble beaches where its grazing can remove vegetation and result in the erosion of sediment, converting soft-bottom shores to rocky habitats (Clay 1961; Bertness 1984; Lubchenco 1986). It is a competitor with native snails and is prey for introduced crabs and many native predators, including carnivorous snails, crabs, fishes, and birds (Dexter 1947; Brenchley and Carlton 1983). It is also a host to many parasites, including digenean trematodes which infect fishes and birds in later stages of their life cycle. One of these parasites, Cryptocotyle lingua, was introduced to North America along with L. littorea (Stunkard 1930; Blakeslee and Byers 2008; Blakeslee et al. 2008).

Economic Impacts

Fisheries: Littorina littorea is a common food item in European cultures, but is rarely part of American cuisine. However, it is frequently eaten by European, Asian, and African immigrants. In the British Isles, 'winkles' are eaten raw or roasted, and the animal extracted from the shell with a bent pin (Clay 1961). Cohen (2012) found L. littorea for sale at 6 of 11 Asian seafood markets, which were visited in San Francisco and Los Angeles. We have no estimate of the economic value of this snail's harvest on North American shores.

Ecological Impacts

Competition: Littorina littorea competes with native shore molluscs for food and living space on Northwest Atlantic shores. Manipulative experiments show that L. littorea can decrease the growth rate of the native L. saxatilis (Rock Periwinkle) and Notoacmea testudinalis (Tortoiseshell Limpet). Field observations and experiments indicate that the Eastern Mudsnail (Ilyanassa obsoleta) emigrates to avoid large concentrations of L. littorea (Brenchley and Carlton 1983). After the Littorina invasion, I. obsoleta abandoned pilings and cobble beaches, and became rare in peat habitats of marshes, habitats where it was formerly abundant. It is now largely restricted to soft-mud habitats not used by L. littorea (Brenchley and Carlton 1983). One of the mechanisms of this competition is interference – L. littorea will climb onto the shells of I. obsoleta and graze algae growing on the shell, causing the mudsnail to twist to dislodge the periwinkle, interfering with the mudsnail's feeding (Brenchley and Carlton 1983).

Herbivory: Littorina littorea is now the most abundant large invertebrate herbivore on the Northwest Atlantic Coast from the Gulf of St. Lawrence to New Jersey. Addition and removal experiments suggest a strong impact on ephemeral green algae (Ulva spp.) in the intertidal, but it also grazes germlings of larger canopy-forming algae, such as Chondrus crispus and Fucus spp. (Lubchenco 1978; Lubchenco and Menge 1978; Lubchenko 1980; Lubchenco 1983; Petraitis 1983; Bertness 1984; Lubchenco 1986). Impacts of L. littorea were greatest at sheltered sites, since L. littorea was rare or absent at sites with heavy wave action (Lubchenco 1978). Littorina littorea also grazes the leaves and rhizomes of the marsh grass Spartina alterniflora, reducing its growth and reproduction on a cobble beach in Rhode Island (Bertness 1984). In a marsh environment in Maine, grazing of L. littorea on S. alterniflora affected the grass' growth only in stressful habitats (poor drainage, long tidal flooding) but did not affect biomass in more optimal habitats. Grazing by Littorina littorea may control the recruitment of introduced Codium fragile ssp. fragile in the upper intertidal, where this alga is also stressed by ultraviolet, freezing, and desiccation. Grazing has less effect in the lower intertidal, where Codium suffers less damage and grows at a faster rate (Scheibling et al. 2008).

Habitat Change: Grazing by Littorina littorea in the rocky intertidal helps to benefit canopy-forming seaweeds such as Chondrus crispus and Fucus spp. by removing ephemeral algal competitors. The canopy-forming seaweeds provide shelter for many species of intertidal invertebrates (Lubchenco and Menge 1978; Lubchenco 1980; Lubchenco 1983; Petratis 1983; Bertness 1984). Grazing by L. littorea had different effects on the Rock Barnacle Semibalanus balanoides (in Long Island sound NY) depending on snail density. Moderate densities favored barnacle settlement by removing algae, but high densities of snails removed large numbers of barnacle cyprids (Petraitis 1983).

Removal experiments on a cobble beach in Mount Hope Bay, Rhode Island indicate that herbivory by Littorina littorea has major effects on habitat structure by removing algae and Spartina alterniflora (Smooth Cordgrass), preventing the accumulation of sediment and removing habitat for soft-bottom fauna (Bertness 1984). Bertness suggests that the Littorina invasion may have resulted in the reduction of marsh habitat and other soft-bottom habitats (Bertness 1984). In marsh environments in the Wells National Estuarine Research Reserve, Maine removal experiments showed that in stressed sites (long inundation, poor drainage), L. littorea reduced the growth of S. alterniflora, but had no effect at a benign site (Tyrrell et al. 2008). Grazing by this introduced snail could reduce the biomass of Spartina at stressed sites, increasing rates of erosion, particularly as a response to sea-level rise (Tyrrell et al. 2008). The extent and magnitude of L. littorea impacts on Spartina may vary among habitats.

Predation: Littorina littorea is a major predator on the egg capsules of I. obsoleta (Eastern Mudsnail) in Barnstable Harbor, Massachusetts (Brenchley 1982). The effect of this predation on the abundance of I. obsoleta is not clear.

Food/Prey: The high biomass of L. littorea at many sites may be benefiting two invading crab species, the Green Crab (Carcinus maenas) and Asian Shore Crab (Hemigrapsus sanguineus), as Littorina littorea is a frequent food item for both species (H. s., Gerard et al. 1999; C. m., Vermeij 1982; Ropes 1989). A wide range of native predators feed on the Common Periwinkle, including predatory snails, crabs, fishes, shorebirds, gulls, and crows (Dexter 1947; Peterson 1979; Ellis et al. 2007).

Parasite/Predator Vector: Littorina littorea has served as a probable vector for one parasite introduced from the coast of Europe, the digenean trematode, Cryptocotyle lingua. This parasite infects Littorina spp. from eggs scattered in the feces of birds and mammals and then develops into the next stage, rediae, which consume some of the snail's tissues, before developing into swimming cercariae, which leave the snail and infect the tissues of fishes, as metacercariae. When the fishes are eaten by birds or mammals, the trematodes mature and shed eggs (Stunkard 1930; Sindermann and Farrin 1962). Genetic analyses of L. littorea and C. lingua from many sites on both sides of the Atlantic, indicates that both the snail and trematode have undergone a similar reduction in genetic diversity and were introduced at roughly the same time (Blakeslee and Byers 2008; Blakeslee et al. 2008). The early stages of the trematode also occur in the native periwinkles L. obtusata and L. saxatilis (Blakeslee and Byers 2008; Blakeslee et al. 2008). In fishes, the cercariae can cause unpleasant pigment spots in fish flesh, while heavy infections can kill young Atlantic Herring (Clupea harengus) (Sindermann and Farrin 1962). In birds ('terns', probably Common Terns, Sterna sterna) and dogs fed infected fish, localized intestinal damage occurred, followed by the development of immunity to infection (Willey and Stunkard 1942). The introduction of L. littorea thus appears to have introduced a parasite that has infected three trophic levels of the Northwest Atlantic food web.

Regional Impacts

NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactHerbivory
Grazing by Littorina littorea may control the recruitment of Codium fragile ssp. fragile in the upper intertidal, where this alga is also stressed by ultraviolet, freezing, and dessication. Grazing by L. littorina has less effect in the lower intertidal, where Codium suffers less damage and grows at a faster rate (Scheibling et al. 2008)
NA-ET3Cape Cod to Cape HatterasEcological ImpactHerbivory
Removal experiments in Mount Hope Bay RI indicate that herbivory by Littorina littorea removes leafy algae such as Ulva spp. on a cobble beach, resulting in the growth of an algal canopy (Bertness 1984). In Port Jefferson Harbor NY, Long Island Sound, grazing by L. littorea also reduced abundance of Ulva, though dense patches were grazed only at high densities, and the alga managed to persist in irregularities in the rocks (Petraitis 1983). In Mount Hope Bay, grazing by L. littorea also reduced the growth and reproduction of the marsh grass Spartina alterniflora (Smooth Cordgrass) by consuming rhizomes of the plant (Bertness 1984).
NA-ET3Cape Cod to Cape HatterasEcological ImpactHabitat Change
Removal experiments in Mount Hope Bay, RI indicate that herbivory by Littorina littorea has major effects on habitat structure, by removing algae and Spartina, preventing the accumulation of sediment, and removing habitat for soft-bottom fauna (Bertness 1984). Bertness suggests that the Littorina invasion may have resulted in the reduction of marsh habitat and other soft-bottom habitats (Bertness 1984). Grazing by L. littorea had different effects on the Rock Barnacle Semibalanus balanoides (in Long Island Sound, NY) depending on snail density. Moderate densities favored barnacle settlement by removing algae, but high densities of snails removed large numbers of barnacle cyprids (Petraitis 1983).
NA-ET2Bay of Fundy to Cape CodEcological ImpactHerbivory
Littorina littorea is the most abundant herbivorous intertidal invertebrate on rocky shores in the Gulf of Maine (Bertness et al. 2002; Bertnes et al. 2004). Addition and removal experiments indicated that L. littorea did not graze significantly on the dominant alga, Chondrus crispus, when it was already established, but did affect algal succession by grazing on emphemeral algae (e.g. Ulva, Enteromorpha spp. which compete with Chondrus, damping variation in Chondrus abundance (Lubchenco 1978; Lubchenco and Menge 1978). The periwinkles graze the germlings of the larger brown seaweed Fucus spp., but also benefit Fucus by removing competitive ephemerals and epibiotic algae (Lubchenko 1980; Lubchenco 1983; Lubchenco 1986) Impacts of L. littorea were greatest at sheltered sites (Canoe Beach Cove, Nahant, MA and Grindstone Neck ME), since L. littorea was rare or absent at sites with heavy wave action (Lubchenco and Menge 1978). In the tidal Damariscotta River estuary, heavy grazing by L. littorea limited growth of most algal species,in areas with slow current flow, leading to the domination of Ascophyllum nodosum (Bertness et al. 2002). In marsh environments in the Wells National Estuarine Research Reserve, ME, removal experiments showed that in a stressed site (long inundation, poor drainage), L. littorea reduced the growth of the marsh grass Spartina alterniflora (Smooth Cordgrass), but had no effect at a benign site (Tyrrell et al. 2008).
NA-ET2Bay of Fundy to Cape CodEcological ImpactHabitat Change
Grazing by Littorina littorea helps to benefit canopy-forming seaweeds such as Chondrus crispus spp. and Fucus spp. by removing ephemeral alga competitors. The canopy forming seaweeds provide shelter for many species of intertidal invertebrates (Lubchenco and Menge 1978; Lubchenco 1980; Lubchenco 1983). In marsh environments in the Wells National Estuarine Research Reserve, ME, removal experiments showed that in a stressed site (long inundation, poor drainage), Littorina reduced the growth of the marsh grass Spartina alterniflora (Smooth Cordgrass), but had no effect at a benign site (Tyrrell et al. 2008). Grazing by this introduced snail could reduce the biomass of Spartina at stressed sites, increasing rates of erosion, particularly as a response to sea-level rise (Tyrrell et al. 2008)
N170Massachusetts BayEcological ImpactHerbivory
Littorina littorea is the most abundant herbivorous intertidal invertebrate on rocky shores in the Gulf of Maine. Manipulative experiments indicated that L. littorea did not graze significantly on the dominant alga, Chondrus crispus, when it was already established, but did affect algal succession by grazing on emphemeral algae (e.g. Ulva, Enteromorpha spp. which compete with Chondrus, damping variation in Chondrus abundance; Lubchenco 1978; Lubchenco and Menge 1978). The periwinkles graze the germlings of the larger brown seaweed Fucus spp., but also benefit Fucus by removing competitive ephemerals and epibiotic algae (Lubchenko 1980; Lubchenco 1983; Lubchenco 1986).  Impacts of L. littorea were greatest at a sheltered sites of Canoe Beach Cove, Nahant, MA, since L. littorea was rare or absent at sites with heavy wave action (Lubchenco and Menge 1978).
N036_CDA_N036 (Maine Coastal)Ecological ImpactHerbivory
Littorina littorea is the most abundant herbivorous intertidal invertebrate on rocky shores in the Gulf of Maine. Manipulative experiments indicated that L. littorea did not graze significantly on the dominant alga, Chondrus crispus, when it was already established, but did affect algal succession by grazing on emphemeral algae (e.g. Ulva spp. which compete with Chondrus, damping variation in Chondrus abundance; Lubchenco 1978; Lubchenco and Menge 1978). The periwinkles graze the germlings of the larger brown seaweed Fucus spp., but also benefit Fucus by removing competitive ephemerals and epibiotic algae (Lubchenko 1980; Lubchenco 1983; Lubchenco 1986). Impacts of L. littorea were greatest at sheltered sites, since L. littorea was rare or absent at sites with heavy wave action (Lubchenco 1978). Moderate abundances and impacts of L. littorea were found at Grindstone Neck (near Schoodic Point), Maine (Lubchenco and Menge 1978; Lubchenco 1980; Lubchenco 1983).
N050Penobscot BayEcological ImpactHerbivory
Littorina littorea is the most abundant herbivorous intertidal invertebrate on rocky shores in the Gulf of Maine. Removal experiments indicated that L. littorea did not graze significantly on the dominant alga, Chondrus crispus, when it was already established, but did affect algal succession by grazing on emphemeral algae which compete with Chondrus, damping variation in Chondrus abundance. Impacts of L. littorea were greatest at sheltered sites such as Little Brewster Island, ME, since L. littorea was rare or absent at sites with heavy wave action (Lubchenco 1978).
N170Massachusetts BayEcological ImpactHabitat Change
Grazing by Littorina littorea helps to benefit canopy-forming seaweeds such as Chondrus crispus spp. and Fucus spp. by removing ephemeral alga competitors. The canopy forming seaweeds provide shelter for many species of intertidal invertebrates. Canoe Beach Cove, Nahant, MA was one experimental site where these impacts were observed (Lubchenco and Menge 1978; Lubchenco 1980; Lubchenco 1983).
N036_CDA_N036 (Maine Coastal)Ecological ImpactHabitat Change
Grazing by Littorina littorea helps to benefit canopy-forming seaweeds such as Chondrus crispus spp. and Fucus spp. by removing ephemeral alga competitors. The canopy forming seaweeds provide shelter for many species of intertidal invertebrates (Lubchenco and Menge 1978; Lubchenco 1980; Lubchenco 1983). Moderate abundances and impacts of L. littorea were found at Grindstone Neck (near Schoodic Point), Maine (Lubchenco and Menge 1978; Lubchenco 1980; Lubchenco 1983).
M020Narragansett BayEcological ImpactHerbivory
Removal experiments in Mount Hope Bay, RI, indicate that herbivory by Littorina littorea removes leafy algae such as Ulva spp. on a cobble beach, resulting in the growth of an algal canopy. Grazing by L. littorea also reduced the growth and reproduction of the marsh grass Spartina alterniflora (Smooth Cordgrass) by consuming rhizomes of the plant (Bertness 1984).
M020Narragansett BayEcological ImpactHabitat Change
Removal experiments in Mount Hope Bay, RI, indicate that herbivory by Littorina littorea has major effects on habitat structure, by removing algae and Spartina, preventing the accumulation of sediment, and removing habitat available for soft-bottom fauna (Bertness 1984). Bertness suggests that the Littorina invasion may have resulted in the reduction of marsh habitat and other soft-sediment habitats (Bertness 1984).
N120Wells BayEcological ImpactHerbivory
In marsh environments in the Wells National Estuarine Research Reserve, ME, removal experiments showed that in a stressed site (long inundation, poor drainage), Littorina reduced the growth of the marsh grass Spartina alterniflora (Smooth Cordgrass), but had no effect at a benign site (Tyrrell et al. 2008).
N120Wells BayEcological ImpactHabitat Change
In marsh environments in the Wells National Estuarine Research Reserve, ME, removal experiments showed that in a stressed site (long inundation, poor drainage), Littorina reduced the growth of the marsh grass Spartina alterniflora (Smooth Cordgrass), but had no effect at a benign site (Tyrrell et al. 2008). Grazing by this introduced snail could reduce the biomass of Spartina at stressed sites, increasing rates of erosion, particularly as a response to sea-level rise (Tyrrell et al. 2008)
NA-ET2Bay of Fundy to Cape CodEcological ImpactPredation
Littorina littorea is a major predator on the egg capsules of the snail Ilyanassa obsoleta (Eastern Mudsnail) in Barnstable Harbor MA (Brenchley 1984). The effect of this predation on abundance of I. obsoleta is not clear.
N180Cape Cod BayEcological ImpactPredation
Littorina littorea is a major predator on the egg capsules of the snail Ilyanassa obsoleta (Eastern Mudsnail) in Barnstable Harbor, MA (Brenchley 1984). The effect of this predation on the abundance of I. obsoleta is not clear.
M010Buzzards BayEcological ImpactCompetition
In the Woods Hole area, from 1900 to 1930, Littorina littorea was observed to gradually exclude Ilyanassa obsoleta from cobble shores and wood pilings, and increasingly invading marshes, largely confining I. obsoleta to mudflats (Dimon 1905; Clench 1930, cited by Brenchley and Carlton 1983). However, experimental studies of the mechanisms of competition were not conducted in this region, to our knowledge. In experimental cage manipulations at Nobska Point and Gansett Beach, Woods Hole, MA L. littorea depressed the growth of L. saxatilis (Rough Periwinkle) in the lower intertidal zone, probably due to food competition. However, this effect was not seen in the upper intertidal zone, where L. saxatilis is more abundant (Yamada and Mansour 1987).
NA-ET3Cape Cod to Cape HatterasEcological ImpactCompetition
In the Woods Hole area, from 1900 to 1930, Littorina littorea was observed to gradually exclude Ilyanass obsoleta (Eastern Mudsnail), from cobble shores and wood pilings, and increasingly invading marshes, largely confining I. obsoleta to mudflats (Dimon 1905; Clench 1930, cited by Brenchley and Carlton 1983). However, experimental studies of the mechanisms of competition were not conducted in this region, to our knowledge. In Mount Hope Bay RI, I. obsoleta was observed to emigrate from sites where large numbers of L. littorea had been added (Bertness, unpublished, cited by Brenchley and Carlton 1983). In experimental cage manipulations near Woods Hole MA, L. littorea depressed the growth of L. saxatilis (Rough Periwinkle), probably due to food competition. However, this effect was not seen in the upper interidal zone, where L. saxatilis is more abundant (Yamada and Mansour 1987).
NA-ET2Bay of Fundy to Cape CodEcological ImpactCompetition
In Barnstable Harbor MA, Ilyanassa obsoleta (Eastern Mudsnail) leave areas of mudflat habitat when large numbers of L. littorea are added. This avoidance behavior may be due in part to the tendency of L. littorea to climb onto the shells of I. obsoleta and graze on the shell surface (Brenchley and Carlton 1982). Removal experiments (on Swans Island ME) indicated that growth, weight gain and survival of the native limpet Notoacmea testudinalis was reduced in the presence of L. littorea (Petraitis 1989).
N180Cape Cod BayEcological ImpactCompetition
In Barnstable Harbor MA, Ilyanassa obsoleta (Eastern Mudsnail) leave areas of mudflat habitat when large numbers of L. littorea are added. This avoidance behavior may be due in part to the tendency of L. littorea to climb onto the shells of I. obsoleta and graze on the shell surface (Brenchley and Carlton 1982).
M020Narragansett BayEcological ImpactCompetition
In Mount Hope Bay, RI, I. obsoleta was observed to emigrate from sites where large numbers of L. littorea had been added (Bertness, unpublished, cited by Brenchley and Carlton 1983).
N045_CDA_N045 (Maine Coastal)Ecological ImpactCompetition
Removal experiments (on Swans Island, ME) indicated that growth, weight gain and survival of the native limpet Notoacmea testudinalis was reduced in the presence of L. littorea (Petraitis 1989).
M040Long Island SoundEcological ImpactHerbivory
In Port Jefferson Harbor, NY, Long Island Sound, grazing by L. littorea also reduced abundance of Ulva, though dense patches were grazed only at high densities and the alga managed to persist in irregularities in the rocks (Petraitis 1983).
M040Long Island SoundEcological ImpactHabitat Change
Grazing by L. littorea had different effects on the Rock Barnacle Semibalanus balanoides (in Long Island Sound, NY) depending on snail density. Moderate densities favored barnacle settlement by removing algae, but high densities of snails removed large numbers of barnacle cyprids (Petraitis 1983).
NA-ET3Cape Cod to Cape HatterasEcological ImpactFood/Prey
The high biomasses of L. littorea may have benefited two invading crab species, the Green Crab (Carcinus maenas and Asian Shore Crab (Hemigrapsus sanguineus). Littorina littorea is a frequent food item for both species (H. s., Gerard et al.1999; C. m., Vermeij 1982; Ropes 1989).
NA-ET2Bay of Fundy to Cape CodEcological ImpactFood/Prey
The high biomasses of L. littorea may have benefited two invading crab species, the Green Crab (Carcinus maenas) and Asian Shore Crab (Hemigrapsus sanguineus). Littorina littorea was a frequent food item for both species (H. s., Gerard et al.1999; C. m., Rops 1989). On the Isles of Shoals ME, L. littorea were a major prey for Jonah Crabs (Cancer borealis), Green Crabs, and Dogwhelks (Nucella lapillus) (Ellis et al. 2007).
N070Damariscotta RiverEcological ImpactHerbivory
In the tidal Damariscotta River estuary, heavy grazing by L. littorea limited growth of most algal species, in areas with slow current flow, leading to the domination of Ascophyllum nodosum (Bertness et al. 2002). On exposed Pemaquid Point, exclusion of L. littorea by cages from cleared areas of rocky shore led to much more rapid regrowth of algae than from control areas (Bertness et al. 2004).
N135_CDA_N135 (Piscataqua-Salmon Falls)Ecological ImpactFood/Prey
On the Isles of Shoals, ME, L. littorea were a major prey for Jonah Crabs (Cancer borealis), Green Crabs (Caricnus maenas), and Dogwhelks (Nucella lapillus) (Ellis et al. 2007).
NA-S3NoneEcological ImpactParasite/Predator Vector
The introduction of Littorina littorea to the Northwest Atlantic is believed to have also introduced the digenean trematode Cryptocotyle lingua. Common Periwinkles collected at 6 sites in the Gulf of St. Lawrence, were infected with C. lingua (Blakeslee and Byers 2008). The rediae (first parasitic stages) have infected not only L. littorea, but also native populations of L. saxatilis (Rough Periwinkle) and L. obtusata (Smooth Periwinkle) (Blakeslee and Byers 2008; Blakeslee et al. 2008). The trematodes cause extensive damage to the host's digestive and reproductive systems (Wood et al. 2009). The rediae of C. lingua metamorphose into cercariae, which have a swimming tail and infect fishes (Stunkard 1930; Sindermann et al. 1962). When infected fish are eaten by birds or mammals, the metacercariae grow in the final host's digestive tract and and reproduce (Stunkard 1930; Sindermann et al. 1962). Thus, the introduction of the Common Periwinkle has added a new parasite to 3 trophic levels.
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactParasite/Predator Vector
The introduction of Littorina littorea to the Northwest Atlantic is believed to have also introduced the digenean trematode Cryptocotyle lingua. Common Periwinkles, collected at 4 sites on the Atlantic coast of Maritime Canada, were infected with C. lingua (Blakeslee and Byers 2008). The rediae (first parasitic stages) have infected not only L. littorea, but also native populations of L. saxatilis (Rough Periwinkle) and L. obtusata (Smooth Periwinkle) (Blakeslee and Byers 2008; Blakeslee et al. 2008). The trematodes cause extensive damage to the host's digestive and reproductive systems (Wood et al. 2009). The rediae of C. lingua metamorphose into cercariae, which have a swimming tail and infect fishes (Stunkard 1930; Sindermann et al. 1962). When infected fish are eaten by birds or mammals, the metacercariae grow in the final host's digestive tract and and reproduce (Stunkard 1930; Sindermann et al. 1962). Thus, the introduction of the Common Periwinkle has added a new parasite to 3 trophic levels.
NA-ET2Bay of Fundy to Cape CodEcological ImpactParasite/Predator Vector
The introduction of Littorina littorea to the Northwest Atlantic is believed to have also introduced the digenean trematode Cryptocotyle lingua. Common Periwinkles, collected at 11 sites in the Gulf of Maine, were infected with C. lingua (Pohley et al. 1976; Blakeslee and Byers 2008). The rediae (first parasitic stages) have infected not only L. littorea, but also native populations of L. saxatilis (Rough Periwinkle) and L. obtusata (Smooth Periwinkle) (Pohley 1976; Blakeslee and Byers 2008; Blakeslee et al. 2008). The trematodes cause extensive damage to the host's digestive and reproductive systems (Wood et al. 2009). The rediae of C. lingua metamorphose into cercariae, which have a swimming tail and infect fishes (Stunkard 1930; Sindermann et al. 1962). When infected fish are eaten by birds or mammals, the metacercariae grow in the final host's digestive tract and and reproduce (Stunkard 1930; Sindermann et al. 1962). Thus, the introduction of the Common Periwinkle has added a new parasite to 3 trophic levels.
NA-ET3Cape Cod to Cape HatterasEcological ImpactParasite/Predator Vector
The introduction of Littorina littorea to the Northwest Atlantic is believed to have also introduced the digenean trematode Cryptocotyle lingua. Common Periwinkles, collected at 7 sites from Massachusetts, were infected with C. lingua (Stunkard 1930; Pohley 1976; Blakeslee and Byers 2008). The rediae (first parasitic stages) have infected not only L. littorea, but also native populations of L. saxatilis (Rough Periwinkle) and L. obtusata (Smooth Periwinkle) (Pohley 1976; Blakeslee and Byers 2008; Blakeslee et al. 2008). The trematodes cause extensive damage to the host's digestive and reproductive systems (Wood et al. 2009). The rediae of C. lingua metamorphose into cercariae, which have a swimming tail and infect fishes (Stunkard 1930; Sindermann et al. 1962). When infected fish are eaten by birds or mammals, the metacercariae grow in the final host's digestive tract and reproduce (Stunkard 1930; Sindermann et al. 1962). Thus, the introduction of the Common Periwinkle has added a new parasite to 3 trophic levels.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET2 Bay of Fundy to Cape Cod 1861 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1875 Def Estab
AR-V None 0 Native Estab
AR-III None 0 Native Estab
NEA-II None 0 Native Estab
B-I None 0 Native Estab
B-II None 0 Native Estab
B-III None 0 Native Estab
B-IV None 0 Native Estab
NEA-III None 0 Native Estab
NEA-IV None 0 Native Estab
NEA-V None 0 Native Estab
NA-S3 None 1840 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 1857 Def Estab
MED-III None 1982 Def Unk
NEP-III Alaskan panhandle to N. of Puget Sound 1937 Def Failed
M010 Buzzards Bay 1875 Def Estab
M020 Narragansett Bay 1880 Def Estab
M040 Long Island Sound 1879 Def Estab
M090 Delaware Bay 1928 Def Estab
M060 Hudson River/Raritan Bay 1888 Def Estab
M080 New Jersey Inland Bays 1892 Def Estab
N130 Great Bay 1980 Def Estab
NEP-VI Pt. Conception to Southern Baja California 2002 Def Failed
P050 San Pedro Bay 2002 Def Failed
P290 Puget Sound 1937 Def Failed
NEP-V Northern California to Mid Channel Islands 1968 Def Unk
NEP-IV Puget Sound to Northern California 1943 Def Failed
P135 _CDA_P135 (Mad-Redwood) 1943 Def Failed
P090 San Francisco Bay 1968 Def Unk
P040 Newport Bay 1975 Def Failed
M110 Maryland Inland Bays 1959 Def Estab
M120 Chincoteague Bay 1971 Def Estab
M128 _CDA_M128 (Eastern Lower Delmarva) 1982 Def Estab
M100 Delaware Inland Bays 1952 Def Estab
N195 _CDA_N195 (Cape Cod) 1875 Def Estab
N190 Waquoit Bay 1996 Def Estab
N140 Hampton Harbor 1887 Def Estab
N180 Cape Cod Bay 1872 Def Estab
N170 Massachusetts Bay 1872 Def Estab
N125 _CDA_N125 (Piscataqua-Salmon Falls) 1880 Def Estab
M065 _CDA_M065 (Middle Delaware-Mongaup-Broadhead) 1970 Def Estab
N060 Muscongus Bay 1880 Def Estab
N050 Penobscot Bay 1897 Def Estab
N045 _CDA_N045 (Maine Coastal) 1880 Def Estab
N040 Blue Hill Bay 1971 Def Estab
N036 _CDA_N036 (Maine Coastal) 1880 Def Estab
N010 Passamaquoddy Bay 1862 Def Estab
M070 Barnegat Bay 1894 Def Estab
N080 Sheepscot Bay 1880 Def Estab
M050 Great South Bay 1937 Def Estab
M036 _CDA_M036 (Southern Long Island) 1880 Def Estab
M030 Gardiners Bay 1953 Def Estab
M026 _CDA_M026 (Pawcatuck-Wood) 1897 Def Estab
N070 Damariscotta River 1971 Def Estab
N116 _CDA_N116 (Piscataqua-Salmon Falls) 1887 Def Estab
N110 Saco Bay 1873 Def Estab
N100 Casco Bay 1873 Def Estab
N185 _CDA_N185 (Cape Cod) 1901 Def Estab
NA-S2 None 1882 Def Estab
P093 _CDA_P093 (San Pablo Bay) 1976 Def Failed
N020 Englishman/Machias Bay 1976 Def Estab
N030 Narraguagus Bay 1880 Def Estab
N065 _CDA_N065 (St. George-Sheepscot) 0 Def Estab
N160 Plum Island Sound 1880 Def Estab
N165 _CDA_N165 (Charles) 1878 Def Estab
N175 _CDA_N175 (Cape Cod) 1880 Def Estab
N120 Wells Bay 1880 Def Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 2004 Def Estab
AR-IV None 2015 Def Unk

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
5458 USNM 24618 None 9999-01-01 None Def 45.0000 -63.0000
5459 USNM 34326 1882 9999-01-01 None Def 54.0000 -60.0000
5460 USNM 74548 None 9999-01-01 Dublin Bay Native 53.3000 -6.1000
5463 USNM 123326 None 9999-01-01 Baddeck Def 46.1000 -60.8000
5466 USNM 162726 None 9999-01-01 Anticosti Island Def 49.5000 -63.0000
5467 USNM 181894 None 9999-01-01 Newfoundland Def 52.0000 -56.0000
5468 USNM 185519 None 9999-01-01 Cornwall, Falmouth Native 50.1000 -5.1000
5469 USNM 185520 1846 1846-08-01 Yorkshire, Scarborough Native 54.3000 -0.4000
5470 USNM 185534 None 9999-01-01 Killiney, Near Dublin Native 53.2000 -6.1000
5471 USNM 185535 1869 9999-01-01 Killybego Native 54.6000 -8.4000
5472 USNM 185543 None 9999-01-01 Essex, Southend On Sea Native 51.8000 0.5000
5473 USNM 185544 1870 1870-09-21 Yorkshire, Scarborough Native 54.3000 -0.4000
5474 USNM 185545 None 9999-01-01 Waldringfield Native 52.0000 1.3000
5475 USNM 185546 1863 9999-01-01 Filey Native 52.6000 1.7000
5476 USNM 185547 1868 9999-01-01 Suffolk, Shottisham Creek Native 52.0000 1.4000
5477 USNM 185561 None 9999-01-01 Charente-Maritime, La Rochelle Native 46.2000 -1.0000
5478 USNM 185861 None 9999-01-01 Cornwall, Lands End Native 50.1000 -5.8000
5479 USNM 185884 None 9999-01-01 Yorkshire, Bridlington Native 54.1000 -0.2000
5480 USNM 186072 None 9999-01-01 Frenchman's Bay Def 44.4000 -68.2000
5481 USNM 193387 1880 1880-10-01 Ten Pound Island, Gloucester Harbor Def 42.6000 -70.7000
5482 USNM 224849 None 9999-01-01 Casco Bay Def 43.7000 -70.0000
5483 USNM 224850 None 9999-01-01 Manchester Def 42.6000 -70.8000
5484 USNM 224851 None 9999-01-01 Nahant Def 42.4000 -70.9000
5485 USNM 305489 None 9999-01-01 Kent, Folkstone Native 51.1000 1.2000
5486 USNM 305521 None 9999-01-01 Channel Islands, Jersey Island Native 49.2000 -2.1000
5490 USNM 408698 1970 9999-01-01 Boston, Castle Island Def 42.4000 -71.1000
5491 USNM 408725 1970 9999-01-01 Gloucester, South East Harbor Def 42.7000 -70.7000
5492 USNM 420428 1933 1933-08-01 Brewster, Marshy Sod Def 41.8000 -70.1000
5493 USNM 431150 1927 2027-06-18 Point St. Charles, Near Matarnek River, On Beach Def 50.2000 -65.8000
5494 USNM 435397 1970 9999-01-01 Campobello Def 44.9000 -66.9000
5495 USNM 435399 1970 9999-01-01 Eastport Def 44.9000 -67.0000
5496 USNM 435400 1970 9999-01-01 Duxbury Def 42.0000 -70.7000
5498 USNM 435413 1912 1912-01-01 Rockland Def 44.1000 -69.1000
5499 USNM 450195 2006 2006-05-01 Cape Norman Def 51.6000 -55.9000
5500 USNM 450196 1914 2014-06-17 Cohasset Def 42.2000 -70.8000
5501 USNM 488283 1970 9999-01-01 Thomaston Def 44.1000 -69.2000
5502 USNM 543175 1970 9999-01-01 Boothbay Harbor Def 43.9000 -69.6000
5503 USNM 590064 1970 9999-01-01 Beverly Def 42.6000 -70.8000
5504 USNM 590448 1948 1948-07-04 Nantucket Island, South Beach Def 41.3000 -70.0000
5505 USNM 592331 1947 1947-10-28 Placentia Def 47.2433 -53.9672
5506 USNM 593135 1898 1898-08-01 Grand Manan Island, Flaggs Cove Def 44.7000 -66.8000
5507 USNM 595270 1950 9999-01-01 Mount Hope Bay, Fall River Def 41.7000 -71.1000
5508 USNM 596930 1951 1951-05-07 Ballybunion Native 52.4000 -9.9000
5509 USNM 600984 1949 1949-06-30 Labrador, Red Bay Def 51.7000 -56.4000
5510 USNM 600995 1949 1949-07-07 St. Anthony Def 51.4000 -55.6000
5511 USNM 603087 1951 1951-06-28 Sjaelland, Jaegerspris Native 55.9000 12.0000
5512 USNM 611704 1970 9999-01-01 Montauk Def 41.0000 -72.0000
5513 USNM 617372 1937 1937-01-01 Long Island, Atlantic Beach Def 40.6000 -73.7000
5514 USNM 633647 1970 9999-01-01 Kieler Bucht Native 54.5833 10.4333
5515 USNM 657040 1959 1959-07-26 Mersay River Def 44.0000 -64.7000
5516 USNM 663278 1953 1953-08-02 Summerside, Def 46.4000 -63.8000
5517 USNM 663931 1970 9999-01-01 Long Island, Sheepshead Bay Def 40.6000 -73.9000
5518 USNM 664567 1970 9999-01-01 Isle Of Man, Port Lewaigue, Ramsey Native 54.2300 -4.5700
5519 USNM 664589 1960 1960-06-25 St. Martins Def 45.4000 -65.5000
5520 USNM 664590 1960 1960-06-21 St. Andrews Def 45.1000 -67.0000
5521 USNM 664591 1960 1960-07-06 North Boundary Queensland, Outlet Of Tidal Creek Def 44.6333 -64.0158
5522 USNM 664592 1960 1960-07-02 Digby Neck, Whale River, Whale Cove Def 44.5000 -66.1000
5523 USNM 664593 1960 1960-07-01 Harbourville, 60 Mile NE Of Digby Def 45.1000 -64.8000
5524 USNM 664594 1960 1960-06-23 Alma Def 45.6000 -65.0000
5525 USNM 664602 1951 1951-07-01 Yorkshire, Scarborough Native 54.3000 -0.4000
5526 USNM 709281 1971 1971-03-20 Chincoteague Island, Def 37.9000 -75.4000
5527 USNM 710427 1975 9999-01-01 North Channel, Firth Of Clyde, Girvan Native 55.2500 -4.8500
5528 USNM 714062 1970 9999-01-01 Glamorgan, Porth Kerry Native 51.8000 -3.9000
5529 USNM 714066 1970 9999-01-01 Suffolk, Deben, Debben Native 52.2000 1.0000
5530 USNM 714135 1952 9999-01-01 Lynn Def 42.5000 -71.0000
5531 USNM 715599 1961 1961-11-18 Saugatuck River, Beach Just E Of River Mouth Def 41.1000 -73.4000
5532 USNM 822723 1976 1976-06-17 Milford Def 41.2000 -73.1000
5533 USNM 1034787 1960 1960-04-30 Barnstable Def 41.7000 -70.3000
5535 MCZ 132925 1941 1941-08-08 York Beach Def 43.1714 -70.6094
5536 MCZ 133081 None 9999-01-01 Swampscott Def 42.4709 -70.9176
5537 MCZ 133168 None 9999-01-01 Gloucester, Bass Rocks Def 42.6158 -70.6625
5538 MCZ 144725 None 9999-01-01 Westbrook Def 41.2853 -72.4481
5539 MCZ 163255 1932 9999-01-01 Perry Def 44.9750 -67.0764
5540 MCZ 163256 1932 9999-01-01 Hampton Beach Def 42.9072 -70.8125
5541 MCZ 163257 1932 9999-01-01 Tiverton Def 41.6258 -71.2139
5542 MCZ 174570 None 9999-01-01 Block Island Def 41.1917 -71.5750
5543 MCZ 176393 1946 1946-06-25 Grand Manan, Grand Harbor, Anchorage Beach Def 44.7000 -66.8000
5544 MCZ 176459 1944 1944-07-01 Marblehead, Cliff St., Beach nr. Yacht Club Def 42.5000 -70.8583
5545 MCZ 177528 1946 1946-05-01 Walcheren, WestKapelle Native 51.5000 3.5833
5546 MCZ 182297 None 9999-01-01 Marshfield, Brant Rock Def 42.0917 -70.7061
5547 MCZ 182309 None 9999-01-01 Cape Cod, Provincetown Harbor Def 42.0417 -70.1667
5548 MCZ 182310 None 9999-01-01 Cape Cod, Bourne, Wing's Neck Def 41.7411 -70.5994
5549 MCZ 182311 None 9999-01-01 Cape Cod, Bourne, Monument Beach Def 41.7153 -70.6153
5550 MCZ 183673 1931 2008-08-31 N. Kattegat, Hirtsholmen Id Native 57.0000 11.0000
5551 MCZ 183790 None 9999-01-01 Kvaroy in Luroy, opposite warehouse Native 66.4833 12.9500
5552 MCZ 192628 None 9999-01-01 Deer Island Def 44.9833 -66.9833
5553 MCZ 192670 None 9999-01-01 Cape Breton Island, North Ingonish Def 46.1667 -60.7500
5554 MCZ 192881 1948 1948-07-04 Nantucket Island, South Beach Def 41.2833 -70.0833
5555 MCZ 192882 1948 1948-08-22 Tuckernuck Island Def 41.3000 -70.2583
5556 MCZ 193580 None 9999-01-01 Cotes du Nord, Brehec Native 48.7167 -2.9500
5557 MCZ 193857 1951 1951-07-01 Kittery, Sea Point Def 43.0881 -70.7367
5558 MCZ 196971 None 9999-01-01 Ostend Native 51.2167 2.9167
5559 MCZ 198037 1952 9999-01-01 Indian River Inlet Opening Def 38.6097 -75.0581
5560 MCZ 199230 None 9999-01-01 4 miles E of Durham, Cedar Pt. Def 43.1303 -70.8528
5562 MCZ 201361 1955 1955-06-15 Old Silver Beach, N of Falmouth on Buzzard's Bay Def 41.6267 -70.6367
5563 MCZ 211047 1956 1956-06-17 Sagamore Beach, Sta. 1838 Def 41.7983 -70.5294
5564 MCZ 211135 1956 1956-07-01 South Dartmouth, Sta 1839 Def 41.5917 -70.9417
5565 MCZ 214405 None 9999-01-01 Chatham Def 41.6819 -69.9603
5566 MCZ 215422 1957 1957-08-24 Yarmouth Def 43.8333 -66.1167
5567 MCZ 219664 1958 1958-08-21 Menai Bridge Native 53.2167 -4.1500
5568 MCZ 219689 1958 1958-08-22 Millport Native 55.7500 -4.9500
5569 MCZ 219834 1956 1956-12-01 Orient Beach State Park Def 41.1300 -72.2661
5570 MCZ 219933 None 9999-01-01 Long Island, Sea Cliff Def 40.8489 -73.6453
5571 MCZ 219963 1956 1956-08-01 S. coast Brittany, 12 mi. S. of Quimper, Beg-Meil Native 48.0000 -4.1000
5572 MCZ 223541 1918 2018-06-20 Bristol, Mouth of Warren River Def 41.7042 -71.2972
5573 MCZ 223542 None 9999-01-01 Wickford Def 41.5739 -71.4619
5574 MCZ 223543 1914 2014-06-18 Narragansett Pier Def 41.4322 -71.4569
5575 MCZ 223545 None 9999-01-01 E Gloucester, Brier Neck Def 42.6250 -70.6250
5576 MCZ 223547 1906 2006-05-31 Marblehead, Castle Rock Def 42.5000 -70.8583
5577 MCZ 223549 None 9999-01-01 Cabbage Island Def 43.8419 -69.6075
5578 MCZ 223551 1904 2008-07-04 Revere Def 42.4083 -71.0125
5579 MCZ 223552 1890 1890-08-01 West Manchester Def 42.5667 -70.7833
5580 MCZ 229329 1960 1960-06-23 Alma Def 45.6000 -64.9500
5581 MCZ 229332 1960 1960-06-23 St. Martins Def 45.3500 -65.5333
5582 MCZ 229355 1960 1960-07-03 Lawrencetown Def 44.6333 -63.3500
5583 MCZ 229356 1960 1960-06-21 St. Andrews Def 45.0667 -67.0333
5584 MCZ 229414 1960 1960-07-02 Digby Neck Def 44.4500 -66.1333
5585 MCZ 235001 1961 1961-12-01 Glamorgan, Sully Native 51.6667 -3.6667
5586 MCZ 235002 1961 1961-12-01 Nash Point, Marcross Beach Native 51.4000 -3.5500
5588 MCZ 236890 None 9999-01-01 Oyster Bay Def 40.9217 -73.5058
5589 MCZ 241023 1955 1955-08-30 Long Island, Shinnecock Bay Def 40.8714 -72.4703
5590 MCZ 249253 None 9999-01-01 Dorset, Swanage Native 50.6000 -1.9667
5591 MCZ 249663 1963 1963-08-01 North Trescott, Crowe's neck Def 44.8594 -67.1322
5592 MCZ 253258 1963 1963-07-01 Indian Point Def 44.8333 -66.9500
5593 MCZ 256592 1965 1965-08-01 Blue Hill, Salt Pond Def 44.4008 -68.5753
5594 MCZ 271587 1968 1968-05-01 Near St. Anthony, Chemeillere Bay Def 51.3833 -55.6000
5595 MCZ 271589 1968 1968-06-01 White Bay, Harry's Harbor Def 50.0000 -56.5833
5596 MCZ 316515 1977 1977-09-01 Chamberlain Def 43.8919 -69.4778
5598 MCZ 343651 1971 1971-03-21 Trenton, NE of Trenton Bridge, in mudflats Def 44.4300 -68.3669
5599 MCZ 343654 1971 1971-01-31 Bass Harbor, State Ferry Peir Def 44.2339 -68.3528
5600 MCZ 343656 1971 1971-03-31 New Harbor, Pemaquid Point Light Def 43.8742 -69.4925
5601 MCZ 343658 1970 1970-12-31 Swans Island, Fir Point, state ferry pier Def 44.1583 -68.4300
5602 MCZ 343659 1971 1971-03-31 Prospect Hill Def 43.3500 -70.4703
5603 MCZ 343665 1970 1970-11-08 Thompson Island Def 44.4270 -68.3639
5604 MCZ 343692 1971 1971-06-16 Milbridge Def 44.5353 -67.8814
5605 MCZ 343694 1971 1971-04-23 Lubec, W. Quoddy Head, Coast Guard Station Def 44.8606 -66.9847
5606 MCZ 343887 1971 1971-03-25 Somesville, Somes Harbor Def 44.3611 -68.3286
5607 MCZ 358368 None 9999-01-01 Parker River Def 42.7533 -70.8139
5610 USNM 152573 None 9999-01-01 Murman Coast, Ara Native 69.3667 32.8000
5611 USNM 185516 1863 9999-01-01 Shetland Islands, Balta Sound Native 60.7333 -0.7667
5612 USNM 185517 1847 9999-01-01 Shetland Islands, Stornoway Native 58.2167 -6.3667
5613 USNM 185527 1868 9999-01-01 Shetland Islands, North Mauin Native 60.5000 -1.0000
5614 USNM 185554 1863 1863-04-01 Akershus, Oslo Native 59.9167 10.7500
5615 USNM 185556 None 9999-01-01 Akershus, Drobak Native 59.6633 10.6311
5617 USNM 224847 1970 9999-01-01 Finnmark, Vadso Native 70.0833 29.7667
5618 USNM 664368 1952 9999-01-01 North Holland, En Helden Native 52.5833 4.9167
5619 USNM 664369 1970 9999-01-01 Cantabrico, Coruna, Vigo, Playa De Marin Native 42.8500 -8.4333
5620 USNM 664599 1951 1951-06-01 North Frisian Islands, Sylt Island, Morsum Native 54.8333 8.2000
5621 USNM 666895 1966 1966-05-05 Zeeland, Zuid Bereland Native 51.8500 6.0333
5622 USNM 714160 1920 2020-08-21 Luroy, Kvardy Native 66.4167 12.8500
5623 USNM 714172 1920 2020-08-26 Melfjord, Telnes Native 68.6833 14.8667
5931 Pohley 1976 1975 1975-01-01 Roque Bluffs Def 44.6129 -67.4797
5939 Alexander 1947 1894 1894-01-01 Point Pleasant Def 40.0832 -74.0682
5940 U.S. National Museum of Natural History 2007 None 9999-01-01 None Def 40.1021 -74.0332
5941 U.S. National Museum of Natural History 2007 None 9999-01-01 Barnegat Light/ Def 39.7576 -74.1063
5945 Ford 1892, cited by Bequaet 1943, 1982 1982-01-01 Atlantic City Def 39.3643 -74.4229
5946 U.S. National Museum of Natural History 2007 1953 1953-01-01 Ocean City Def 39.2776 -74.5746
5947 Alexander 1947); Wells 1965 1946 1946-01-01 Intracoastal Waterway jetties, Cape May/ Def 38.9668 -74.9630
5948 Kraueter 1965 1970 1970-01-01 Army Pier, Cape Henlopen Def 38.8032 -75.0946
5949 U.S. National Museum of Natural History 2007 1980 1980-01-01 Lewes Def 38.7929 -75.1580
5950 Museum of Comparative Zoology 2008 1952 1952-01-01 Bethany Beach Def 38.6098 -75.0577
5951 Wells 1965 1959 1958-01-01 Ocean City Def 38.3365 -75.0849
5952 Krauter 1974 1971 1971-01-01 Chincoteague Def 37.9357 -75.3683
5953 Vermeij 1982 1982 1982-01-01 Wachapreague/ Def 39.0220 -74.9064
5958 Reid 1996 1996 1996-01-01 Rugen Native 54.4167 13.4000
5959 Reid 1996 1996 1996-01-01 Bornholm Native 55.1667 15.0000
5960 Reid 1996) 1996 1996-01-01 Stavanger Native 58.9667 5.7500
5961 Reid 1996 1996 1996-01-01 Bergen Native 60.3911 5.3247
5962 Reid 1996 None 9999-01-01 Trondheim Native 63.4167 10.4167
5963 Reid 1996 1996 1996-01-01 Trondheim Native 63.4167 10.4167
5964 Reid 1996 1996 1996-01-01 Steinkjer Native 64.0167 11.5000
5965 Reid 1996 None 9999-01-01 Kandalaksha Native 67.1500 32.4167
5966 Reid 1996 None 9999-01-01 Nordkapp Native 71.1678 25.8025
5967 Reid 1996) None 9999-01-01 Grotoy Native 70.1667 18.8500
5968 Reid 1996 None 9999-01-01 Zakhrebetnoye Native 69.0275 36.4275
5969 Reid 1996 None 9999-01-01 Iokanga Native 68.0531 39.5131
5970 Reid 1996 None 9999-01-01 Solovetskiye Ostrova Native 65.1167 35.8833
5971 Reid 1996 None 9999-01-01 Bourgneuf Native 47.0500 -1.9500
5972 Reid 1996 None 9999-01-01 La Rochelle Native 47.2667 -1.3833
5973 Reid 1996 None 9999-01-01 Arcachon Native 44.6500 -1.1667
5974 Reid 1996 None 9999-01-01 Biarritz Native 43.4833 -1.5667
5976 Reid 1996 None 9999-01-01 Carino Native 43.7500 -7.8667
5977 Reid 1996 None 9999-01-01 Esposende Native 41.5333 -8.7833
5978 Reid 1996 None 9999-01-01 Cascais Native 38.7000 -9.4167
5979 Reid 1996 None 9999-01-01 Sines Native 37.9500 -8.8667
5980 Reid 1996 None 9999-01-01 Faro Native 37.0167 -7.9333

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