Invasion History

First Non-native North American Tidal Record: 1993
First Non-native West Coast Tidal Record: 1993
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Littorina saxatilis has an extensive native distribution in the Atlantic, stretching from the Arctic Ocean (up to 80°N) south to the Mid-Atlantic States (New Jersey-Virginia) and the Iberian Peninsula (Reid 1996). Isolated southern populations in the Azores, Canary Islands, and the Mediterranean Sea (Tunisia) could represent remnants of Pleistocene populations, although fossil evidence is lacking. Natural transport by birds or rafting is also possible. In the Northeast Atlantic, L. saxatilis has colonized the isolated islet of Rockall, between Britain and Iceland (Johannesson 1988). Johannesson (1988) suggests that the ovoviparous life history has adapted this species to colonizing remote sites, since a single fertilized female rafted on driftwood or seaweed, or carried by a ship, can start a population. Alternatively, these southern records could represent early introductions by shipping. On the East coast of North America, the construction of stone jetties and riprap may have favored the extension of L. saxatilis’ range to the Atlantic coast of Virginia, near the mouth of Chesapeake Bay (Counts and Bashore 1991). The type population in Venice Lagoon, Italy, from which the species was first described in 1792, is probably introduced (Panova et al. 2011), as are occurrences in four estuaries on the coasts of South Africa and Namibia (Mead et al. 2011b). The most recent invasion is in San Francisco Bay, where this snail was first collected in 1993 (Carlton and Cohen 1998).

North American Invasion History:

Invasion History on the West Coast:

Littorina saxatilis has been introduced to the West Coast of North America, in San Francisco Bay, California. In 1993, this periwinkle was collected in rocky debris along the shore at Emeryville Marina, on central San Francisco Bay, by James T. Carlton (Carlton and Cohen 1998). In 1996, a second population was found at Taney Marina Beach, Coast Guard Island, on the Oakland estuary (Carlton and Cohen 1998). A research group, led by Whitman Miller of the Smithsonian Environmental Research Center, began a series of surveys in 2002 in order to evaluate the possibility of eradication. They found that L. saxatilis was already sufficiently widespread so that eradication was not feasible. Several populations were found on South San Francisco Bay, including one at the Port of Redwood City (Miller et al. 2004) and in Brisbane Lagoon (Cohen et al. 2005). The probable vector for introduction of this periwinkle was imported baitworms (Neanthes virens) wrapped in seaweed (Ascophyllum nodusum) from Maine and Maritime Canada. The seaweed packing material is usually discarded along the shore, and often contains living marine invertebrates, including L. littorea, L. saxatilis, and L. obtusata (Carlton and Cohen 1998; Miller et al. 2004).

Invasion History Elsewhere in the World:

Littorina saxatilis was first described from Venice Lagoon, Italy in 1792 (Reid 1996). This population has been considered to be one of several cryptogenic remnant populations in Pleistocene refuges (Reid 1996). However, genetic analysis of Venetian specimens shows low genetic diversity and similarity to British and Scandinavian populations, strongly supporting introduced status (Panova et al. 2011). Other isolated populations, in the Gulf of Gabés, Tunisia and in the Azores and Canary Islands, could be Pleistocene remnants, or natural colonists (Johannesson 1988; Reid 1996), but to our knowledge, have not been studied genetically.

Isolated populations of L. saxatilis occur in South Africa and Namibia. It was first reported in 1963 from Langebaan Lagoon, South Africa (Robinson et al. 2004; Robinson et al. 2005; Mead et al. 2011b). Fossils believed to date to about 40,000 years BP, and postglacial fossils, are known from the vicinity of Cape Town (Johannesson 1988; Reid 1996). However, dates have not been determined by radiological methods. As Mead et al. (2011b) have noted, if the fossil populations had survived to the present, this periwinkle would probably be much more widespread on the South African coast.


Description

Littorina saxatilis is a medium-sized marine snail, with a turban-shaped shell, having a moderately elevated, pointed spire. The shell is dextrally coiled and adult shells have 4-5 convex whorls with well-defined sutures. The sculpture of the shell is variable, but consists of many poorly-defined, or as few as five, more definite spiral ribs. The color is highly variable, from drab-gray to yellow-brown or dark brown, and sometimes white, yellow, orange, red, brown, or black. Some individuals, especially young snails, are banded or spotted with yellow and black. Mature snails can range from 1.2 to 25.8 mm, but are more usually 6-13 mm. Development is direct, with young developing in a brood pouch and emerging as tiny snails, about 0.45 to 0.75 mm. Description from: Abbott 1974; Morris 1975; and Reid 1996.

The taxonomy of this snail is one of the most confused of any mollusk, including 28 synonymous names and 31 varieties, resulting from its high morphological variability (Reid 1996). At least two very similar species, L. arcana and L. compressa, are known from European waters, but have not been reported from North American waters (Reid 1996). The occurrence of numerous ecotypes and divergent populations probably stems from brooding reproduction and the limited dispersal ability of this snail (Reid 1996; Mäkinen et al. 2008; Doellman et al. 2011).


Taxonomy

Taxonomic Tree

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

Synonyms

Littorina castanea (Deshayes, 1843)
Littorina davidus (Roding, 1798)
Littorina groenlandica (Menke, 1830)
Littorina jugosa (Montagu, 1803)
Littorina neglecta (Bean, 1844)
Littorina obligatus (Say, 1822)
Littorina rudis (Lyell, 1835)
Littorina rudis var. groenlandica (Menke, 1830)
Littorina rudis var. saxatilis (Harmer, 1921)
Littorina rudis var. tenebrosa (Jeffreys, 1865)
Littorina saxatilis (Brusina, 1870)
Littorina saxatilis rudis (Dautzenberg and Fischer, 1912)
Littorina saxatilis tenebrosa (Dautzenberg and Fischer, 1912)
Littorina tenebrosa (Gould, 1841)
Littorina vestita (Say, 1822)
Turbo rudis (Maton, 1797)
Turbo saxatilis (Olivi, 1792)
Turbo tenebrosus (Montagu, 1803)

Potentially Misidentified Species

Littorina arcana
Hannaford Ellis 1978; Known from the British Isles, Brittany, Normandy (France) and Norway. The shell is morphologically identical to L. saxatilis, but reproduction is oviparous and anatomical details differ (Reid 1996).

Littorina compressa
Jeffreys 1865 (=L. nigrolineata); Known from the British Isles, Brittany (France), and Norway. The shell is strongly ribbed, reproduction is oviparous, and the radular and soft anatomical details differ from L. saxatilis (Reid 1996).

Littorina sitkana
Some erroneous reports of L. saxatilis from the Pacific Ocean have been due to confusion with this northeastern Pacific species (Reid 1996).

Ecology

General:

Littorina saxatilis occurs mostly on mid-to-upper shore rocky coasts, but is also found in lagoons and marshes, usually on hard substrates. Sexes are separate and fertilization is internal. This snail can mature at sizes as small as 1-2 mm. In a Quebec population, L. saxatilis matured at about 3.2-3.5 mm (Pardo and Johnson 2005). Larval development is ovoviviparous, with the young developing in a brood pouch and emerging at 0.4-0.75 mm, with a shell of 1.6-2.1 whorls. The number of embryos varies with size of the female, from 10 in a female with a shell length of 2 mm to 750-900 embryos in the largest shells. Breeding occurs year-round, but is reduced in late summer and winter (Reid 1996).

Littorina saxatilis inhabits a wide range of habitats. On the most exposed coasts, they are usually confined to sheltered crevices, but can also occur in the spray zone, above normal tidelines. More usually, they are found from the mid-tide to mean high-water of spring tides, but they also occur in subtidal regions of brackish lagoons and marshes, associated with solid substrate such as stones, shells, and the roots of Spartina. As with L. littorea, occurrence in the subtidal zone is limited by predation by crabs, fishes, and other predators (Reid 1996). In a Quebec population, the smaller snails tend to be concentrated in the upper intertidal, while larger specimens were more frequently found in the lower zone (Pardo and Johnson 2005). As an upper-intertidal snail, the Rough Periwinkle is tolerant to a wide range of air and water temperatures and also exposure to low salinity, especially over short durations (Reid 1996; Sokalova and Portner 2003). However, its range is probably limited by high temperatures in southern Europe, where it is largely confined to lagoonal habitats and the West Atlantic coast south of New York Harbor, where it is restricted to man-made rocky habitats (Reid 1996; Counts and Bashore 1991; Prezant et al. 2002).

Littorina saxatilis grazes unselectively on diatoms, cyanobacteria, and filamentous and leafy macroalgae (Reid 1996). Littorina saxatilis and L. littorea are potential competitors, and overlap in the intertidal zone, but L. saxatilis ranges higher up on the shore (Dexter 1945; Clay 1961). Predators include crabs, including Carcinus maenas (Green Crab), fishes, and shorebirds (Reid 1996). This snail also hosts a variety of parasites. Fourteen trematode species were found in L. saxatilis sampled at 29 East Coast sites from Newfoundland to Connecticut, but only 4 of those trematodes are known from its introduced range in San Francisco Bay (Blakeslee et al. 2012).

Food:

Macroalgae; microalgae

Consumers:

Fish; Shorebirds; land mammals

Competitors:

Littorina littorea

Trophic Status:

Herbivore

Herb

Habitats

General HabitatMarinas & DocksNone
General HabitatRockyNone
General HabitatSalt-brackish marshNone
General HabitatCanalsNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Salinity RangeMesohaline5-18 PSU
Tidal RangeMid IntertidalNone
Tidal RangeHigh IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-20In air at low tide (Sokalova and Portner 2006)
Maximum Temperature (ºC)35In air at low tide, tolerates short term exposures up to 47 C (Sokalova and Portner 2006)
Minimum Salinity (‰)5Field observations for estuarine populations (Reid 1996) A marine White Sea population had more than 30% mortality at 14 ppt (Sokalova 2000). Snails from two Swedish Skaggerak (high-salinity) populations ceased activity at 16 ppt. Snails from an exposed coastal region, found in upper tidepools, survived longer in fresh water (17 days) than those from a sheltered region, occurring lower on the shore (Sundell 1985). Todd (1964) found that survival at low salinities varied with temperature, with lower limits of 8 PSU at 5 C, but 16 at 15 C.
Minimum Length (mm)1.2Minimum size of mature snail (Reid 1996)
Maximum Length (mm)25.8More usually 6-13 mm (Abbott 1974; Morris 1975; Reid 1996)
Broad Temperature RangeNonePolar-Warm Temperate
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Impacts of Littorina saxatilis have not been studied in San Francisco Bay or in other invaded areas. However, since L. saxatilis is larger than native Northeast Pacific periwinkles and survives under a very wide range of environmental conditions, it is potentially capable of competing with and displacing native snails from California to Alaska. The native periwinkles L. scutulata, L. plena and L. keanae were absent in the immediate vicinity of introduced L. saxatilis populations in San Francisco Bay (Carlton and Cohen 1998), suggesting possible competition or behavioral exclusion. If L. saxatilis extends its range north of San Francisco Bay competition with the morphologically similar species, L. subrotundata and L. sitkana is possible (Carlton and Cohen 1998). Modeling based on the present range of L. saxatilis, and ocean conditions, indicates that it could potentially colonize the entire West Coast from California to the Arctic Ocean (de Rivera et al. 2011). If this snail develops large populations, then impacts from increased grazing on intertidal algal communities are possible (Carlton and Cohen 1998).

Partly because of potential impacts, but also based on this snail's limited introduced range and dispersal, an attempt was made to eradicate introduced populations in San Francisco Bay (Smithsonian Environmental Research Center 2004). However, in the 9+ years since its introduction, L. saxatilis had spread over enough area of the coastline to render eradication impractical (Ruiz et al., unpublished data).

Regional Impacts

NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
The native periwinkles L. scutulata, L. plena and L. keenae were absent in the vicinity of introduced L. saxatilis populations in San Francisco Bay (Carlton and Cohen 1993). However, the mechanisms for this apparent exclusion have not been studied, to our knowledge.
P090San Francisco BayEcological ImpactCompetition
The native periwinkles L. scutulata, L. plena and L. keenae were absent in the vicinity of introduced L. saxatilis populations in San Francisco Bay (Carlton and Cohen 1993). However, the mechanisms for this apparent exclusion have not been studied, to our knowledge.

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEA-V None 0 Native Estab
NEA-IV None 0 Native Estab
NEA-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
AR-V None 0 Native Estab
AR-III None 0 Native Estab
AR-IV None 0 Native Estab
NA-P3 None 0 Native Estab
NA-P4 None 0 Native Estab
NA-S2 None 0 Native Estab
NA-S3 None 0 Native Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 0 Native Estab
NA-ET2 Bay of Fundy to Cape Cod 0 Native Estab
NA-ET3 Cape Cod to Cape Hatteras 0 Native Estab
NEA-VI None 0 Crypto Estab
MED-IV None 1792 Crypto Estab
NEP-V Northern California to Mid Channel Islands 1993 Def Estab
WA-IV None 1969 Def Estab
P090 San Francisco Bay 1993 Def Estab
M128 _CDA_M128 (Eastern Lower Delmarva) 0 Native Estab
M120 Chincoteague Bay 1971 Native Estab
M100 Delaware Inland Bays 0 Native Estab
M090 Delaware Bay 0 Native Estab
M080 New Jersey Inland Bays 0 Native Estab
M070 Barnegat Bay 0 Native Estab
M040 Long Island Sound 0 Native Estab
M030 Gardiners Bay 0 Native Estab
M060 Hudson River/Raritan Bay 0 Native Estab
M026 _CDA_M026 (Pawcatuck-Wood) 0 Native Estab
M020 Narragansett Bay 0 Native Estab
M010 Buzzards Bay 0 Native Estab
N185 _CDA_N185 (Cape Cod) 0 Native Estab
N170 Massachusetts Bay 0 Native Estab
N165 _CDA_N165 (Charles) 1880 Native Estab
N180 Cape Cod Bay 0 Native Estab
N195 _CDA_N195 (Cape Cod) 0 Native Estab
N130 Great Bay 0 Native Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 0 Native Estab
N060 Muscongus Bay 0 Native Estab
N100 Casco Bay 0 Native Estab
N010 Passamaquoddy Bay 0 Native Estab
N030 Narraguagus Bay 0 Native Estab
N045 _CDA_N045 (Maine Coastal) 0 Native Estab
N116 _CDA_N116 (Piscataqua-Salmon Falls) 0 Native Estab
N080 Sheepscot Bay 0 Native Estab
N036 _CDA_N036 (Maine Coastal) 0 Native Estab
N050 Penobscot Bay 0 Native Estab
NA-P2 None 0 Native Estab
WA-I None 0 Crypto Estab
MED-VII None 1792 Def Estab
MED-II None 0 Crypto Unk
NA-S1 None 0 Native Estab
WA-V None 1969 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
3684 USNM 225946, US Museum of Natural History 2002 1970 1970-01-01 Smith's Island Native 37.1439 -75.8700
3685 USNM 709282 Museum of Natural History 2002 1971 1971-03-20 None Native 37.9333 -75.3833
3687 USNM 803497 , US Museum of Natural History 2002 1980 1980-09-10 Breakwater Harbor, Roosevelt Inlet Native 38.7922 -75.1150
3688 USNM 516193 , US Museum of Natural History 2002 1924 1924-09-01 Cape May Native 38.9350 -74.9064
3689 USNM 435456 , US Museum of Natural History 2002 1970 1970-01-01 Atlantic City Native 39.3708 -74.4374
3690 US Museum of Natural History 2002 1970 1970-01-01 Keyport Native 40.4331 -74.2000
3691 USNM 336289, US Museum of Natural History 2002 None 9999-01-01 Manhasset Bay, Long Island Sound Native 40.8307 -73.7221
3692 USNM 404656 , US Museum of Natural History 2002 1928 2028-05-20 Orient/ Native 41.1389 -72.3039
3693 USNM709132, US Museum of Natural History 200 1966 1966-07-18 Hunter Island, Bronx, New York City Native 40.8745 -73.7904
3694 USNM 714122, US Museum of Natural History 2002 1950 1950-07-01 None Native 40.8918 -73.4821
3695 USNM 714125 US Museum of Natural History 2002 1953 1953-01-01 New Rochelle Native 40.9114 -73.7828
3696 USNM 714133, US Museum of Natural History 2002 1953 1953-01-01 Shell Beach, Shelter Island/ Native 41.0428 -72.3439
3697 USNM 435455 , US Museum of Natural History 2002 1970 1970-01-01 Bay Ridge, Brooklyn Native 40.6340 -74.0146
3698 USNM 589963 , US Museum of Natural History 2002 1970 1970-01-01 Stonington Native 41.3333 -71.9000
3699 USNM 762417, US Museum of Natural History 2002 1976 1976-01-01 Branford Native 41.2794 -72.8156
3700 USNM 709606, US Museum of Natural History 2002 1976 1976-01-01 Watch Hill Native 41.3000 -71.9500
3701 USNM 665782, US Museum of Natural History 2002 1970 1970-01-01 Point Judith Native 41.3611 -71.4811
3702 USNM 408693, US Museum of Natural History 2002 1970 1970-01-01 Onset Native 41.7417 -70.6583
3703 USNM 435351, US Museum of Natural History 2002 1910 1910-01-01 South Dennis Native 41.6894 -70.1569
3705 U.S. National Museum of Natural History 2002 None 9999-01-01 Nahant Native 42.4167 -72.9167
3706 U.S. National Museum of Natural History 2002 1880 1880-10-01 Ten Pound Island, Glouceste Native 42.6000 -70.6667
3707 U.S. National Museum of Natural History 2002 None 9999-01-01 Chelsea Beach Native 42.4000 -71.0333
3708 U.S. National Museum of Natural History 2002 1913 1913-01-01 Duxbury Native 40.0333 -70.6667
3709 U.S. National Museum of Natural History 2002 1970 1970-01-01 Provincetown Native 42.0583 -70.1792
3710 U.S. National Museum of Natural History 2002 1946 1946-01-01 West Jetty, Nantucket Island Native 41.2667 -70.0500
3711 U.S. National Museum of Natural History 2002 1951 1951-01-01 Martha's Vineyard, Edgartown Native 41.3500 -70.6667
3712 U.S. National Museum of Natural History 2002 1970 1970-01-01 Newington Native 43.1001 -70.8337
3713 U.S. National Museum of Natural History 2002 1970 1970-01-01 Hampton Beach Native 42.9072 -70.8125
3714 U.S. National Museum of Natural History 2002 1970 1970-01-01 St. Georges Banks Native 44.0165 -69.1989
3715 U.S. National Museum of Natural History 2002 1970 1970-01-01 None Native 43.6667 -70.0000
3716 U.S. National Museum of Natural History 2002 1893 1893-08-09 Eastport, Shacksfords Head Native 44.9000 -66.0000
3717 U.S. National Museum of Natural History 2002 1970 1970-01-01 Five Islands/ Native 44.5670 -67.7992
3718 U.S. National Museum of Natural History 2002 1970 1970-01-01 Isle au Haut Native 44.0753 -68.6339
3719 U.S. National Museum of Natural History 2002 1907 1907-01-01 Kennebunkport Native 43.3833 -70.5500
3720 U.S. National Museum of Natural History 2002 1970 1970-01-01 Boothbay Native 43.8764 -69.6342
3721 U.S. National Museum of Natural History 2002 1970 1970-01-01 Bar Harbor Native 44.3958 -68.1939
3722 U.S. National Museum of Natural History 2002) 1981 1981-08-27 Moose Point State Park Native 44.4394 -68.9483
3723 U.S. National Museum of Natural History 2020 1893 1893-08-01 Grand Manan Island, Flaggs Cove Native 42.7000 -66.8167
3724 U.S. National Museum of Natural History 2002 1981 1981-08-22 Pocologan Native 45.1333 -66.5833
3725 U.S. National Museum of Natural History 2002 1970 1970-01-01 Arichat Native 45.5108 -61.0153
3726 U.S. National Museum of Natural History 2002 1927 1927-10-01 None Native 45.8000 -64.3825
3727 U.S. National Museum of Natural History 2002 1970 1970-01-01 Peggys Cove Native 44.4931 -63.9167
3728 U.S. National Museum of Natural History 2002 1981 1981-08-25 Bay of Fundy Native 44.6833 -65.7825
3729 U.S. National Museum of Natural History 2002 1981 1981-08-25 Port Maitland Native 43.9819 -66.1469
3730 U.S. National Museum of Natural History 2002 1970 1970-01-01 St. Peters Native 45.6561 -60.8750
3731 U.S. National Museum of Natural History 2002 None 9999-01-01 Beach Between Great Whale River + Richmond Gulf Native 57.0000 -77.0000
3732 U.S. National Museum of Natural History 2002 1929 1929-07-01 Trois Pistoles Native 48.1167 -69.1833
3733 U.S. National Museum of Natural History 2002 1929 1929-07-14 Quebec Native 61.6167 -72.0000
3734 U.S. National Museum of Natural History 2002 1927 1927-01-01 None Native 47.0000 -62.0000
3735 U.S. National Museum of Natural History 2002 1946 1946-01-01 St. Simeon Native 48.0667 -65.5667
3736 U.S. National Museum of Natural History 2002 1946 1946-06-25 None Native 51.4500 -56.9667
3737 U.S. National Museum of Natural History 2002 1970 1970-01-01 Riviere du Loup Native 47.8333 -69.5333
3738 U.S. National Museum of Natural History 2002 1925 1925-01-01 None Native 54.0000 -60.0000
3739 U.S. National Museum of Natural History 2002 1882 1882-01-01 Dead Island, Fox Island, Henley Harbo Native 52.8000 -55.8656
3740 U.S. National Museum of Natural History 2002 1906 1906-01-01 Port Saunders Native 50.6453 -57.3056
3742 U.S. National Museum of Natural History 2002 1908 1908-01-01 Nain Native 56.5500 -61.6833
3743 U.S. National Museum of Natural History 2002 1949 1949-08-09 Hebron Native 58.2000 -62.6158
3744 U.S. National Museum of Natural History 2002 1949 1949-07-28 None Native 55.3000 -59.3333
3745 U.S. National Museum of Natural History 2002 1949 1949-07-24 None Native 53.8667 -57.0000
3746 U.S. National Museum of Natural History 2002 1951 1951-08-02 Hopedale Harbour Native 55.4500 -60.2158
3747 U.S. National Museum of Natural History 2002 1951 1951-07-19 None Native 54.3000 -57.7000
3748 U.S. National Museum of Natural History 2002 1951 1951-07-19 None Native 54.5500 -57.1822
3749 U.S. National Museum of Natural History 2002 1949 1949-06-30 None Native 51.7328 -56.4278
3750 U.S. National Museum of Natural History 2002 1979 1979-07-21 None Native 47.6264 -52.6650
3751 U.S. National Museum of Natural History 2002 None 9999-01-01 Salmonier Native 47.0642 -55.2094
3752 U.S. National Museum of Natural History 2002 1906 1906-05-01 Cape Norman Native 51.6192 -55.8992
3753 U.S. National Museum of Natural History 2002 None 9999-01-01 None Native 62.0000 -70.0000
3754 U.S. National Museum of Natural History 2002 None 9999-01-01 Cape Churchill Native 57.1667 -93.7500
3755 U.S. National Museum of Natural History 2002 1929 1929-10-17 Port Churchill Native 58.7667 -94.1667
3756 U.S. National Museum of Natural History 2002 1933 1933-08-06 5 Mile W Of Manico Point, Southampton Island Native 64.5000 -84.5000
3767 U.S. National Museum of Natural History 2002 None 9999-01-01 Signuia Native 62.9000 -64.6167
3768 U.S. National Museum of Natural History 2002 None 9999-01-01 Baffin Island Native 63.4667 -67.4000
3769 U.S. National Museum of Natural History 2002 1927 1927-01-01 Baffin Island Prb 68.5000 -70.0000
3770 U.S. National Museum of Natural History 2002 1943 1943-08-30 Baffin Island Native 63.4667 -67.3833
3771 U.S. National Museum of Natural History 2002 1942 1942-08-12 Near Sylvia Grinnel River, Baffin Island Native 63.4667 -67.3833
3920 U.S. National Museum of Natural History 2002 1959 1959-08-15 Cape Farewell Native 59.7667 -43.9167
3921 U.S. National Museum of Natural History 2002 1874 1875-07-11 Disko Island/ Native 69.7500 -55.5000
3923 U.S. National Museum of Natural History 2002 1875 1875-01-01 Holsteinborg Native 67.0000 -52.0000
3924 U.S. National Museum of Natural History 2002 1970 1970-01-01 Godthab Native 64.1833 -51.7500
3925 U.S. National Museum of Natural History 2002 1970 1970-01-01 None Native 69.7667 -51.3000
3926 U.S. National Museum of Natural History 2002 1931 1931-08-28 Angmagssalik Native 65.5000 -37.5500
3927 U.S. National Museum of Natural History 2002 1920 1920-01-01 None Native 65.3000 -13.8333
3928 U.S. National Museum of Natural History 2002 None 1970-01-01 None Native 65.8333 -15.7167
3929 U. S. Museum of Natural History 2002 None 9999-01-01 None Native 65.5000 38.0000
3930 U. S. Museum of Natural History 2002 1937 1937-01-01 Kola Peninsula Native 69.5000 36.0000
3931 U. S. Museum of Natural History 2002 None 9999-01-01 Finmark Native 70.0000 26.0000
3932 U. S. Museum of Natural History 2002 1970 1970-01-01 None Native 60.1667 6.0000
3933 U. S. Museum of Natural History 2002 1970 1970-01-01 Lofoten Islands/ Native 68.4711 13.8636
3934 U. S. Museum of Natural History 2002 1970 1970-01-01 Vadso, Native 70.0833 29.7667
3935 U. S. Museum of Natural History 2002 1917 1917-07-03 Bjorn Native 66.0833 12.5833
3936 U. S. Museum of Natural History 2002 1917 1917-07-06 Brønnøy Native 65.4667 12.2167
3937 U. S. Museum of Natural History 2002 1970 1970-01-01 None Native 66.5167 13.2500
3949 U.S. National Museum of Natural History 2002 1878 1878-05-01 near Bergen Native 60.5333 5.3000
3950 U.S. National Museum of Natural History 2002 None 1863-04-01 Christiania Native 59.9167 10.7500
3951 U.S. National Museum of Natural History 2002 1970 1970-01-01 Spitzbergen Native 78.0000 15.0000
3952 U.S. National Museum of Natural History 2002 None 9999-01-01 Vallo Native 59.2608 10.4992
3953 U.S. National Museum of Natural History 2002 1970 1970-01-01 None Native 54.5833 10.5833
3954 U.S. National Museum of Natural History 2002 1951 1951-11-01 Sylt Native 54.9000 8.3333
3955 U.S. National Museum of Natural History 2002 1868 1868-01-01 St. Magnus Bay Native 60.4000 -1.5667
3956 U.S. National Museum of Natural History 2002 None 1970-01-01 near the Sea/ Native 51.6333 1.1500
3957 U.S. National Museum of Natural History 2002 1970 1970-01-01 Guernsey Island Native 49.0333 -2.0333
3958 U.S. National Museum of Natural History 2002 None 9999-01-01 None Native 55.6667 -5.0000
3959 U.S. National Museum of Natural History 2002 None 9999-01-01 Portrush Native 55.2167 -6.6500
3960 U.S. National Museum of Natural History 2002 None 9999-01-01 Dunmore Native 52.1514 -6.9861
3961 U.S. Museum of Natural History 2002 1970 1970-01-01 Ryde, Isle of Wight Native 50.7167 -1.1667
3962 U.S. Museum of Natural History 2002 1951 1951-05-07 None Native 52.0581 -9.9378
3963 U.S. Museum of Natural History 2002 1970 1970-01-01 None Native 53.3333 -6.1333
3964 U.S. Museum of Natural History 2002 None 9999-01-01 Penarth Head Native 51.4333 -3.1667
3965 U.S. Museum of Natural History 2002 1905 1905-01-01 Lynmouth, Native 50.4000 -4.2667
3966 U.S. Museum of Natural History 2002 None 9999-01-01 Ballachulish Native 56.6667 -5.1167
3967 U.S. Museum of Natural History 2002 None 9999-01-01 Aberdeen Native 57.1333 -2.1000
3968 U.S. Museum of Natural History 2002 1869 1869-01-01 None Native 54.9167 -8.0000
3969 U.S. Museum of Natural History 2002 None 9999-01-01 Scarborough, Native 54.2833 -0.4333
3970 U.S. Museum of Natural History 2002 1970 1970-01-01 Dinard Native 48.6333 -2.0667
3971 U.S. Museum of Natural History 2002 None 9999-01-01 None Native 47.2167 -1.5500
3972 U.S. Museum of Natural History 2002 1957 1957-08-03 La Coruna Native 43.3667 -8.3833
3973 U.S. Museum of Natural History 2002 None 1947-05-10 Arrecife, Lanzarote Native 28.9500 -13.5330
3974 U.S. Museum of Natural History 2002 None 9999-01-01 Androusa Crypto 37.1167 21.9333
3975 U.S. Museum of Natural History 2002 None 1883-01-01 Lido, Venice Def 45.5833 12.3500
3977 Academy of Natural Sciences of Philadelphia 2004 1973 1973-08-06 Reykjanes Native 63.8333 -22.6833
3978 Johannson 1988 None 9999-01-01 None Crypto 38.5000 -28.0000
3979 Johannson 1988 None 9999-01-01 Rockall Native 57.5969 -13.6886
3980 Johannson 1988 None 9999-01-01 None Native 36.1167 -5.3333
3981 Johannson 1988 None 9999-01-01 Novaya Zemlaya Native 72.0000 53.0000
3982 Dautzberg 1897, cited by Johannsson 1988 1897 1897-01-01 Tunis Crypto 33.8022 9.8533
3983 Gruet 1977 LOOKUP 1977 1977-01-01 Sion sur L'Ocean Native 46.7167 -1.9833
3984 Backeljauet al. 1984LOOKUP 1984 9999-01-01 Store Ekkeroya, Native 70.0833 30.1333
3985 Geertz-Hansen et al. 1993LOOKUP 1993 1993-01-01 Roskilde Native 55.9333 12.0000
3986 Zander et al. 2000 LOOKUP 2000 2000-01-01 Salzhaff ( Native 54.0667 11.5833
3987 Arrontes et al. 2004LOOKUP 1998 1998-01-01 Campiello Native 43.5500 -6.4000
3988 Robinson et al. 2005 None 9999-01-01 Saldanha Def -33.1333 18.0333
3989 Robinson et al. 2005 None 9999-01-01 None Def -34.0333 23.0333
3991 Carlton and Cohen 1998 1996 1996-05-28 Taney Marina Beach, Coast Guard Island Def 37.7822 -122.2489
3992 Miller et al. unpublished 2002 9999-01-01 Port of Redwood City Def 37.5133 -122.2086
4030 Reid 1996 1919 1919-01-01 Cape Barrow Native 68.0167 -110.1667
4031 Reid 1996 1996 9999-01-01 Skinnastadur/ Native 66.0000 -16.5000
4032 Reid 1996 None 9999-01-01 Akureyri Native 65.6667 -18.1000
4033 Reid 1996 None 9999-01-01 Dyrholaey Native 63.4000 -19.1333
4034 Reid 1996 None 9999-01-01 Danskoy Native 79.5000 10.0000
4035 Reid 1996 None 9999-01-01 Kapp Platen Native 80.0000 25.0000
4100 Reid 1996 None 9999-01-01 Jan Mayen Island Native 74.4167 19.0000
4101 Reid 1996 None 9999-01-01 Andenes, Lofoten Islands Native 69.3203 16.1428
4102 Reid 1996 None 9999-01-01 Reine, Lofoten Islands Native 68.0833 13.5167
4103 Reid 1996 None 9999-01-01 Maløy Native 61.9294 5.1192
4104 Reid 1996 None 9999-01-01 Trondheim Native 63.4167 10.4167
4105 Reid 1996 None 9999-01-01 Mandal Native 58.0333 7.4500
4106 Reid 1996 None 9999-01-01 Molde Native 62.7667 7.5000
4107 Reid 1996 None 9999-01-01 Westkapelle Native 51.5333 3.4500
4108 Reid 1996 None 9999-01-01 Mimizan Native 44.2000 -1.2333
4109 Reid 1996 None 9999-01-01 Bilbao Native 43.2500 -2.9667
4110 Reid 1996 None 9999-01-01 Oporto Native 41.1500 -8.6167
4111 Reid 1996 None 9999-01-01 Lisbon Native 38.7167 -9.1333
4112 Reid 1996 None 9999-01-01 Huelva Native 37.2583 -6.9508
4113 Reid 1996 None 9999-01-01 Kandalaksha Native 67.1500 32.4167
6911 Mead et al. 2011b 1969 1969-01-01 Velddriff Def -32.7833 18.1667
28114 Cohen, et al. 2005 (SF Bay Area RAS) 2004 2004-05-27 Coyote Point Marina, San Francisco Bay Def 37.5907 -122.3180
28910 Cohen et al. 2005; Foss 2009 2005 2005-07-06 Coyote Point Def 37.5920 -122.3210
31360 Cohen, et al. 2005 (SF Bay Area RAS) 2004 2004-05-24 Coast Guard Island Marina, San Francisco Bay Def 37.7812 -122.2458
33607 Cohen, et al. 2005 (SF Bay Area RAS) 2004 2004-05-23 Brisbane Lagoon, San Francisco Bay Def 37.6862 -122.3906

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