Invasion History

First Non-native North American Tidal Record: 1987
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1987

General Invasion History:

The encrusting bryozoan Membranipora membranacea was described from the Baltic Sea in Sweden by Linnaeus in 1767. In its native European waters, it occurs from the Barents Sea to the Atlantic coast of Spain, usually growing on macroalgae (Kluge 1975; Schwaninger 1999). It had been previously considered to have a global anti-tropical distribution, in the Northern and Southern Hemispheres, but North Pacific populations have been regarded as separate species by some taxonomists (Dick et al. 2005), while some or all Southern Hemisphere occurrences have been attributed to introductions by shipping (Keough and Ross 1999; Gappa 2000; Griffiths et al. 2009). However, mitochondrial DNA studies indicate that this 'species' consists of three long-separated clades in the North Pacific, and monophyletic clades in the North Atlantic, Southeast Pacific (Chile), Southwest Pacific (Australia, New Zealand), and Southeast Atlantic (South Africa) (Schwaninger 1999; Schwaninger 2008). The South African population has been described as a new species, M. rustica (Florence et al. 2007, cited by Mead et al. 2011b). This species complex may have originated in the North Pacific and colonized the Southern Ocean (Chile, South Africa, Australia, New Zealand) 10-20 million years ago, before reaching the Northeast Atlantic (Schwaninger 2008). The only verified invasion is its introduction from the Northeast Atlantic to the Northwest Atlantic. It was first reported from the Isles of Shoals, New Hampshire in 1987, and now occurs from Long Island Sound to southern Labrador (Berman et al. 1992; Wantanabe et al. 2010; Fisheries and Oceans Canada 2011; USGS Nonindigenous Species Program 2011).

North American Invasion History:

Invasion History on the East Coast:

Membranipora membranacea was first reported from kelp beds around the Isles of Shoals (New-Hampshire-Maine), and from Cape Neddick, Maine, in 1987 (Berman et al. 1992; Lambert et al. 1992). It was first reported south of Cape Cod at Mohegan Bluffs, Block Island, Rhode Island (1990, USGS Nonindigenous Aquatic Species Program 2011). Its southern limit appears to be Long Island Sound (1993, USGS Nonindigenous Aquatic Species Program 2011). Its northward spread has been more extensive. It has colonized Great Bay, Casco Bay, and Penobscot Bay in the Gulf of Maine (Whitlatch and Osman 2000; Pratt and Grason 2007; USGS Nonindigenous Aquatic Species Program 2011), and in 1992 was found on the Atlantic Coast of Nova Scotia, in Mahone and St. Margarets Bay (Saunders and Metaxas 2007). By 2002, it reached the western coast of Newfoundland, on the Gulf of St. Lawrence, and by 2009 was reported from several bays on the south and north coasts of the island. Its current northernmost record is Red Bay, Labrador, on the Straits of Belle Isle (Fisheries and Oceans 2011).


Description

Colonies of the encrusting Membranipora membranacea form an extensive white lacy covering on algae, often on kelp of the genus Laminaria. Colonies on other alga, such as Fucus spp., are smaller. Young colonies are circular, but growth tends to be fastest towards the base of the frond, which is where new growth of the kelp is occurring. This pattern of growth keeps the colony spreading away from the most encrusted areas, and from the areas of the algal frond most likely to be torn off by waves (Hayward and Ryland 1998; Winston and Hayward 2012).

The autozooids of M. membranacea are rectangular, about 0.4 X 0.15 mm in size, with tubercles or short spines at the corners. Tips of these spines are uncalcified. The whole frontal surface is covered by membranes. Each lateral wall has two vertical, uncalcified bands, which permit the colony to adhere to an algal frond, and bend as the alga flexes with the waves. Some zooids may be 'tower zooids', with huge outward projections. The polypide has about 17 tentacles. The cyphonautes larva is planktotrophic, about 0.6 by 0.8 mm in size (description from Hayward and Ryland 1998; Winston and Hayward 2012), illustrated in Newell and Newell (1977).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Bryozoa
Class:   Gymnolaemata
Order:   Cheilostomata
Suborder:   Anasca
Family:   Membraniporidae
Genus:   Membranipora
Species:   membranacea

Synonyms

Flustra membranacea (Linnaeus, 1767)

Potentially Misidentified Species

Membranipora chesapeakensis
Before it was described as a new species, Membranipora chesapeakensis ( Banta et al. 1995) was found growing on Ruppia maritima by Osburn (1944) and was identified as M. membranacea (Osburn 1944; Dudley 1973; Banta et al. 1995).

Membranipora rustica
The South African form of 'M. membranacea' has been described as a new species, M. rustica (Florence 2007; et al. 2011b).

Membranipora serrilamella
Dick et al. (2005) consider M. villosa and M. serrilamella to be two distinct species in the North Pacific. However, Schwaninger (1999; 2008), on the basis of DNA and other studies, considered these species to be inducible morphs of the North Pacific clade of M. membranacea.

Membranipora villosa
Dick et al. (2005) consider M. villosa and M. serilamella to distinct species in the North Pacific. However, Schwaninger (1999; 2008), on the basis of DNA and other studies, considered these species to be inducible morphs of the North Pacific clade of M. membranacea.

Ecology

General:

Life History- Membranipora membranacea is an encrusting, calcified bryozoan, composed of many individual zooids. The zooids feed by extending the ciliated tentacles of the lophophore as a funnel, creating a current, and driving food particles into their mouths. The food is guided along the tentacles and through the pharynx by the cilia. Larger food particles can be moved or captured by flicking or contracting the tentacles (Barnes 1983). Larvae of M. membranacea are planktotrophic, and have a planktonic period of about 24 weeks (Yoshioka et al. 1982). Larvae settle on a substrate, usually an alga, and metamorphose into the first zooid of a colony, an ancestrula (Dudley 1973; Barnes 1983).

Ecology- Membranipora membranacea is known primarily from seaweed beds, especially kelps such as Laminaria and Saccharina spp. but also Agarum clathratum, Desmarestia aculeata, Fucus spp, and the introduced Codium fragile fragile (Hayward and Ryland 1998; Harris and Tyrell 2001). In addition, it occurs on ships' hulls, buoys, and fouling panels (Woods Hole Oceanographic Institution 1952; Dijkstra and Harris 2009; Ruiz et al. unpublished data). It is characteristic of cooler waters, often with active water motion, and penetrates to a limited degree into reduced salinities (Winston 1977; Hayward and Ryland 1998).

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatVessel HullNone
General HabitatGrass BedNone
General HabitatCoarse Woody DebrisNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-1.8Freezing point of seawater, based on range (Saunders and Metaxas 2007)
Maximum Temperature (ºC)26Experimental, acclimated at 18 C, temperature increased at 2 C per day (Menon 1972).
Minimum Salinity (‰)27Field data, Great Bay NH (Blezard 1999)
Minimum Reproductive Temperature8Start of larval settlement, July, Nova Scotia (Saunders and Metaxas 2007)
Maximum Reproductive Temperature21Pacific populations, La Jolla CA (conspecificity uncertain, Yoshioka 1982)
Minimum Duration28Estimated larval duration, field, Pacific (Schwaniger 1999)
Maximum Duration60Estimated larval duration, field, Pacific (Schwaniger 1999)
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Economic Impacts

Fisheries- Membranipora membranacea affects human economies mostly through its effects on cultured large kelps (Saccharina and Laminaria spp.). This bryozoan fouls the surface of kelp fronds, reducing growth, and making the kelp prone to breakage. In waters of Maine and Atlantic Canada, kelp is cultured primarily for industrial chemicals. Bryozoan fouling in Chaleur Bay, Quebec, was reduced by setting out cultures in the fall and harvesting them before heavy larval settlement in July (Gendron et al. 2010). Membranipora membranacea can potentially affect may other fisheries, since kelp beds are prime habitat for lobsters, crabs, sea urchins, and many species of commercial fish. Heavy settlement by this bryozoan reduces kelp survival and biomass, and favors dominance by algae which provide less cover (including the introduced Codium fragile) (Lambert et al. 1992; Harris and Tyrell 2001; Watanabe et al. 2010). However, the effects of its invasion on fisheries yields cannot be easily estimated.

Ecological Impacts

Membranipora membranacea has negative impacts on seaweeds by fouling their surfaces, reducing their rate of photosynthesis, growth, and spore production, and making them more prone to tearing and breakage by water surges during storms. In some cases, M. membranacea invasions have led to great reductions in kelp biomass (Berman et al. 1992; Lambert et al. 1992; Saier and Chapman 2004; Saunders and Metaxas 2008).

Competition- Membranipora membranacea also competes for space with other epibionts, such as other bryozoans (e.g. Electra crustulenta and hydroids Obelia geniculata), and usually overgrows them (Berman et al. 1992; Lambert et al. 1992).

Habitat Change- Membranipora membranacea invasions in the Gulf of Maine, and the Atlantic coast of Nova Scotia have resulted in great reductions of biomass of large kelps (Saccharina and Laminaria spp.), favoring their replacement by smaller algae which are less vulnerable to fouling damage (e.g. Agarum clathratum, Desmarestia aculeata, and the introduced Codium fragile), and reducing habitat cover (Harris and Tyrell 2001; Saunders and Metaxas 2008; Watanabe et al. 2010). Membranipora membranacea has also affected invaded habitats by increasing the breakdown of kelp into detritus, greatly increasing detritus production (Krumhansl and Scheibling 2011).

Food/Prey- Membranipora membranacea has provided a greatly increased food supply for at least one bryozoan predator, the nudibranch Onchidoris muricata, possibly allowing it to reproduce earlier in the season (Pratt and Grason 2007). On a broader scale, this bryozoan may increase both the quantity and quality of detritus, by speeding the breakup of kelp (Krumhansl and Scheibling 2011), and by adding CaCo3 and protein to the kelp eaten by sea urchins (Strongylocentrotus droebachiensis), resulting in richer fecal material, and more potential food for microbes (Sauchyn and Scheibling 2009).

Regional Impacts

NA-ET2Bay of Fundy to Cape CodEcological ImpactCompetition
On the Isles of Shoals (NH-ME) Membranipora membranacea consistently overgrows the bryozoan Electra pilosa and the hydroid Obelia geniculata on kelp fronds, and is rarely overgrown by other epiphytes (Berman et al. 1992). At Cape Neddick, Membranipora membranacea covered extensive areas (22-52%) of the area of kelp (Laminaria saccharum = Saccharina latissima) fronds, potentially decreasing growth and photosynthesis. The bryozoan also largely replaced other kelp epibionts (Lambert et al. 1992).
NA-ET2Bay of Fundy to Cape CodEcological ImpactHabitat Change
Kelps overgrown by M. membranacea are more vulnerable to breaking and tearing during storm surges (Berman et al. 1992). Fouling by Membranipora membranacea killed kelps (Saccharina and Laminaria spp.) in the Cape Cod Canal, Massachusetts, and Cape Neddick, Maine (Lambert et al. 1992). Decreases in the abundance of large kelps (Laminaria spp.) at the Isles of Shoals have contributed to increases in other seaweeds (Agarum clathratum, Desmarestia aculeata, and the introduced Codium fragile fragile), which suffer less damage from colonization by M. membranipora (Harris and Tyrell 2001).
NA-ET2Bay of Fundy to Cape CodEcological ImpactFood/Prey
Membranipora membranacea provides an additional food resource for the nudibranch Onchidoris muricata, possibly allowing it to reproduce earlier in the season (Pratt and Grason 2007; Lambert et al. 2016).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactCompetition
Fouling by M. membranacea killed kelps (Saccharina longicuris = Laminaria spp.) in St. Margaret's Bay by encrusting the fronds and making them more vulnerable to breakage (Saunders and Metaxas 2008) and decreasing spore production (Saier and Chapman 2004). Membranipora membranacea also out-competed the native bryozoan Electra pilosa when growing on kelps, with faster growth rates. However, on rockweeds (Fucus evanescens-native and F. serratus- introduced), E. pilosa grew more rapidly, and frequently out-competed M. membranacea, whose growth rate was reduced on Fucus spp. In the field, M. membranacea is scarce on F. evanescens and absent on F. serratus, while E. pilosa dominates on both species (Yorke and Metaxas 2011).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactHabitat Change
Fouling by M. membranacea results in widespread defoliation of kelps (Laminaria sp., decreasing habitat cover (Saunders and Metaxas 2008; Scheibling and Gagnon 2009). Membranipora mebranacea fouling was positively correlated to an index of summer temperature, while kelp cover was negatively correlated. The mortality of kelp in turn favors the colonization of the kelp beds by the introduced green alga Codium fragile (Scheibling and Gagnon 2006). Bryozoan fouling inhibits grazing on kelp by the snail Lacuna vincta (Banded Chink Shell), resulting in more concentrated grazing on unfouled areas of kelp blades, possibly increasing the risk of the the kelp breaking up. However, no adverse effects on the snail were found (O'Brien et al. 2013). Fouling by M. membranacea greatly increases detrital production in kelp beds (Krumhansl and Scheibling 2011).
N135_CDA_N135 (Piscataqua-Salmon Falls)Ecological ImpactCompetition
In the Isles of Shoals (NH-ME), Membranipora membranacea's colonies overgrew large areas of the fronds of kelp (Laminaria saccharina = Saccharina latissima), potentially decreasing the photosynthesis and growth of the seaweed. In addition, M. membranacea consistently overgrows the bryozoan Electra pilosa and the hydroid Obelia geniculata on kelp fronds, and is rarely overgrown by other epiphytes (Berman et al. 1992). Over time, from 1996-1999, in the Isles of Shoals, M. membranacea increasingly colonized other algal species, Agarum clathratum, Desmarestia aculeata, and the introduced Codium fragile (Harris and Tyrell 2001).
N135_CDA_N135 (Piscataqua-Salmon Falls)Ecological ImpactHabitat Change
Kelps overgrown by M. membranacea are more vulnerable to breaking and tearing during storm surges (Berman et al. 1992). Decreases in the abundance of large kelps (Saccharina = Laminaria spp.) at the Isles of Shoals have contributed to increases in other seaweeds (Agarum clathratum, Desmarestia aculeata, and the introduced Codium fragile fagile, which suffer less damage from colonization by M. membranipora (Harris and Tyrell 2001).
N125_CDA_N125 (Piscataqua-Salmon Falls)Ecological ImpactCompetition
At Cape Neddick, Membranipora membranacea covered extensive areas (22-52%) of the area of kelp (Laminaria saccharina = Saccharina latissima) fronds, potentially decreasing growth and photosynthesis. The bryozoan also largely replaced other kelp epibionts (Lambert et al. 1992).
N125_CDA_N125 (Piscataqua-Salmon Falls)Ecological ImpactHabitat Change
Kelps overgrown by M. membranacea are more vulnerable to breaking and tearing during storm surges (Berman et al. 1992). Fouling by Membranipora membranacea killed kelps (Laminaria spp.) at Cape Neddick, Maine (Lambert et al. 1992).
N125_CDA_N125 (Piscataqua-Salmon Falls)Ecological ImpactFood/Prey
Membranipora membranacea provides an additional food resource for the nudibranch Onchidoris muricata, possibly allowing it to reproduce earlier in the season (Pratt and Grason 2007; Membranipora membranacea provides an additional food resource for the nudibranch Onchidoris muricata, possibly allowing it to reproduce earlier in the season (Pratt and Grason 2007; Lambert et al. 2016).)
N100Casco BayEcological ImpactFood/Prey
Membranipora membranacea provides an additional food resource for the nudibranch Onchidoris muricata, possibly allowing it to reproduce earlier in the season (Pratt and Grason 2007).
N180Cape Cod BayEcological ImpactCompetition
Extensive fouling by M. membranacea led to widespread death of kelp (Saccharina and Laminaria spp.) in the Cape Cod Canal (Lambert et al. 1992).
N180Cape Cod BayEcological ImpactHabitat Change
Extensive fouling by M. membranacea led to widespread death of kelp (Saccharina and Laminaria spp.) in the Cape Cod Canal (Lambert et al. 1992).
NA-S3NoneEconomic ImpactFisheries
Settlement of M. membrancea adversely affected the quality of kelp Saccharina longicruris during aquaculture trails in Gaspe, Quebec. Starting the cultures in the fall, and harvesting the kelp before the settlement of M. membranacea alleviated these problems (Gendron et al. 2010).
NA-ET1Gulf of St. Lawrence to Bay of FundyEcological ImpactFood/Prey
Fouling by M. membranacea greatly increases detrital production in kelp beds (Krumhansl and Scheibling 2011). In addition, kelp encrusted by M. membranacea have a lower absorption efficiency when eaten by sea urchins (Strongylocentrotus droebachiensis), resulting in richer fecal material, and more potential food for microbes (Sauchyn and Scheibling 2009).
N130Great BayEcological ImpactFood/Prey
Membranipora membranacea provides an additional food resource for the nudibranch Onchidoris muricata, possibly allowing it to reproduce earlier in the season (Pratt and Grason 2007; Lambert et al. 2016).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEA-II None 0 Native Estab
NEA-III None 0 Native Estab
NEA-IV None 0 Native Estab
AR-V None 0 Native Estab
NEA-V None 0 Native Estab
NA-ET2 Bay of Fundy to Cape Cod 1987 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 1992 Def Estab
B-II None 0 Native Estab
B-I None 0 Native Estab
SA-I None 0 Crypto Estab
AUS-VIII None 1879 Native Estab
AUS-VII None 0 Native Estab
AUS-X None 0 Native Estab
AR-III None 0 Native Estab
SEP-A' None 0 Native Estab
NA-ET3 Cape Cod to Cape Hatteras 1990 Def Estab
N130 Great Bay 1989 Def Estab
M010 Buzzards Bay 2000 Def Estab
M040 Long Island Sound 1993 Def Estab
M020 Narragansett Bay 2000 Def Estab
SEP-B None 0 Native Estab
NZ-IV None 0 Native Estab
M026 _CDA_M026 (Pawcatuck-Wood) 1990 Def Estab
N050 Penobscot Bay 1998 Def Estab
N070 Damariscotta River 1992 Def Estab
N100 Casco Bay 2004 Def Estab
N110 Saco Bay 1998 Def Estab
N125 _CDA_N125 (Piscataqua-Salmon Falls) 1987 Def Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 1987 Def Estab
N170 Massachusetts Bay 2000 Def Estab
N180 Cape Cod Bay 1992 Def Estab
N140 Hampton Harbor 2007 Def Estab
N120 Wells Bay 2007 Def Estab
N080 Sheepscot Bay 2007 Def Estab
NA-S3 None 2002 Def Estab
NA-S2 None 2009 Def Estab
B-IV None 0 Native Estab
WA-I None 1909 Crypto Unk

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
7902 None None 9999-01-01 None Def 41.1259 -73.2396
7903 USGS Nonindigenous Aquatic Species Program 1990 1990-01-01 Mohegan Bluffs, Block Island Def 41.1518 -71.5567
7904 USGS Nonindigenous Aquatic Species Program 2008 1990 1990-01-01 Quonochontaug Beach Def 41.3318 -71.7217
7905 MacIntyre et al. 2010) 2010 2010-01-01 Fort Adams State Park, Newport Def 41.4782 -71.3356
7906 MIT Sea Grant 2003 2000 2000-01-01 Newport Shipyard, Newport Def 41.4826 -71.3256
7907 MIT Sea Grant 2003 2000 2000-01-01 Tripp Marina Def 41.5168 -71.0856
7908 MIT Sea Grant 2003 2000 2000-01-01 Massachusetts Maritime Academy, Bourne Def 41.7395 -70.6239
7909 MIT Sea Grant 2003 2000 2000-01-01 Sandwich Def 41.7704 -70.5036
7910 Whitlach and Osman 2000 1998 1998-01-01 Plymouth Harbor Def 41.9751 -70.6661
7911 MIT Sea Grant 2003 2000 2000-01-01 Quincy Def 42.2395 -70.9764
7912 MIT Sea Grant 2003) 2000 2000-01-01 Boston Harbor Def 42.3834 -71.0453
7913 Bell et al. 2005 2003 2003-01-01 Boston Harbor Islands Def 42.3301 -70.9284
7914 MIT Sea Grant 2003) 2000 2000-01-01 Salem Harbor Def 42.5220 -70.8823
7915 MIT Sea Grant 2003 2000 2000-01-01 Gloucester Harbor Def 42.5959 -70.6689
7916 MIT Sea Grant 2009 2007 2007-01-01 Hampton River Marina, Hampton Def 42.8995 -70.8209
7917 Berman et al. 1993; Harris and Tyrell 2000 1987 1987-01-01 Isles of Shoals Def 42.9865 -70.6120
7919 USGS Nonindigenous Aquatic Species Program 2008 1989 1989-01-01 near Fox Pt. Def 43.1212 -70.8589
7920 Ruiz et al. unpublished data 2001 2001-01-01 Badgers Island Marina, Portsmouth Def 43.0823 -70.7517
7921 Whitlatch and Osman 2000 1998 1998-01-01 York Harbor Def 43.1306 -70.6312
7922 Berman et al. 1992; Lambert et al. 1992 1987 1987-01-01 Cape Neddick Def 43.1698 -70.6034
7923 Whitlatch and Osman 2000 1998 1998-01-01 Cape Porpoise Def 43.3731 -70.4295
7924 Pratt and Grason 2007 2004 2004-01-01 Bailey Island Def 43.7376 -69.9937
7925 MIT Sea Grant 2008 2007 2007-01-01 Port Harbor Marine, South Portland Def 43.6415 -70.2409
7926 MIT Sea Grant 2008 2007 2007-01-01 Brewer South Freeport Marina Def 43.8198 -70.1095
7927 MIT Sea Grant 2008 2007 2007-01-01 Maine DMR Docks, Boothbay Harbor Def 43.8465 -69.6348
7928 Lambert et al. 1992; MIT Sea Grant 2008 1992 1992-01-01 Darling Maine Center Dock, Wapole Def 43.9401 -69.5737
7929 Whitlatch and Osman 2000 1998 1998-01-01 Castine Harbor Def 44.3829 -68.7989
7930 Whitlatch and Osman 2000 1998 1998-01-01 Belfast Bay Def 44.4281 -69.0020
7931 MIT Sea Grant 2009 2007 2007-01-01 Journey's End Marina, Rockland Def 44.1045 -69.1017
7932 Wantanabe et al. 2010 2007 2007-01-01 Cape Sable Def 43.4070 -65.5050
7933 Watanabe et al. 2009 2007 9999-01-01 Sydney Def 46.1530 -60.1170
7934 Saunders and Metaxas 2007 1992 1992-01-01 St. Margarets Bay Def 44.7730 -63.9530
7935 Saunders and Metaxas 2007 1992 1992-01-01 Mahone Bay Def 44.4540 -64.3530
7936 Watanabe et al. 2009 2007 2007-01-01 Canso Def 45.3660 -60.9900
7937 Watanabe et al. 2009 2007 2007-01-01 None Def 45.0240 -62.2130
7938 Fisheries and Oceans Canada 2011 2010 2010-01-01 St. Marys Bay Def 46.9290 -53.6790
7939 Fisheries and Oceans Canada 2011 2009 2009-01-01 Placentia Def 47.2430 -53.9270
7940 Fisheries and Oceans Canada 2011 2010 2010-01-01 Bay l'Argent Def 47.5503 -54.8714
7941 Fisheries and Oceans Canada 2011 2010 2010-01-01 Harbour Breton Def 47.4670 -55.8440
7942 Gendron et al. 2010) 2006 2006-01-01 Paspebiac/ Def 48.0061 -65.2833
7943 Fisheries and Oceans Canada 2011 2002 2002-01-01 Mouse Island Def 47.6070 -59.1230
7944 Fisheries and Oceans Canada 2011 2002 2002-01-01 None Def 48.7370 -58.9260
7945 Fisheries and Oceans Canada 2011 2009 2009-01-01 Lark Harbour Def 49.1050 -58.4110
7946 Fisheries and Oceans Canada 2011 2009 2009-01-01 Point Riche Def 50.7070 -57.3910
7947 Caines and Gagnon 2012 2009 2009-01-01 Norris Point Def 49.5530 -57.8380
7948 Caines and Gagnon 2012 2008 2008-01-01 Daniels Harbour Def 50.2330 -57.5890
7949 Fisheries and Oceans Canada 2011 2009 2009-01-01 Anchor Point Def 51.3320 -56.7280
7950 Caines and Gagnon 2012 2008 2008-01-01 Green Island Cove Def 51.3790 -56.5750
7951 Caines and Gagnon 2012 2008 2008-01-01 Red Bay Def 51.8050 -56.4000
7953 Fisheries and Ocean Canada 2011 2009 2009-01-01 Witless Bay Def 47.3560 -52.7460
7954 Fisheries and Ocean Canada 2011 2009 2009-01-01 St. Johns Def 47.5675 -52.7072
7955 Fisheries and Ocean Canada 2011) 2009 2009-01-01 Holyrood Def 47.4340 -53.1140
7956 Fisheries and Ocean Canada 2011 2010 2010-01-01 Dover Def 48.9900 -53.9230
7957 Fisheries and Oceans Canada 2011 2009 2009-01-01 Lewisporte Def 49.2520 -55.0390
7958 Swaninger 2008 None 9999-01-01 Ribadeo Native 43.5333 -7.0333
7959 Nielsen and Worsaae 2010 None 9999-01-01 Kristineberg Native 55.5500 13.0667
7960 Schwaniger 1999 None 9999-01-01 Isle of Man Native 54.2500 -4.5000
7961 Schwaninger 2008 None 9999-01-01 Kaldbak, Faroe Islands Native 62.0631 -6.8261
7962 Pascoe et al. 2007 None 9999-01-01 Plymouth Native 50.3714 -4.1424
7964 Gruhl 2008 None 9999-01-01 Concarneau Native 47.8761 -3.9178
7965 Telnes 2008 None 9999-01-01 Trondheim Fjord Native 63.5000 10.4667
7966 Nylund and Pavia 2005 None 9999-01-01 Tjärnö Marine Biological Laboratory Native 58.9333 11.1667
7967 Seed and Harris 1980 None 9999-01-01 Strangford Lough Native 54.4830 -5.5830
7968 Carrada 1973 None 9999-01-01 Ria de Vigo Native 42.2500 -8.7500
7969 Menon 1973 None 9999-01-01 Helgoland Native 54.1825 7.8853
7970 Leloup 1967 None 9999-01-01 Ostend Native 51.2333 2.9167
7971 Thompson et al. 1966 None 9999-01-01 Farne Islands Native 55.6220 1.6280
7972 Brattegard 1966 None 9999-01-01 Hardangerfjord Native 60.1666 6.0000
7973 Kluge 1975 None 9999-01-01 Vardo Native 70.3706 31.1075
7974 Kluge 1975 None 9999-01-01 Kola Inlet Native 69.0833 33.3967
7975 Kluge 1975, None 9999-01-01 Dal'ne Zelenetskaya Inlet Native 69.1100 36.0800
7976 Marzinelli et al. 2011 None 9999-01-01 Balmoral Beach Native -33.8253 151.2464
7977 Schwaninger 2008 None 9999-01-01 Port Phillip Bay Native -39.1500 144.8667
7978 Schwaninger 2008 None 9999-01-01 Noarlunga Jetty, Port Noarlunga Native -35.1500 138.4667
7979 Schwaninger 2008 None 9999-01-01 Point Turton, Adelaide Native -34.9469 137.3531
7980 Schwaninger 2008 None 9999-01-01 Wellington Native -41.2889 174.7772
7981 Schwaninger 2008 None 9999-01-01 Concepcion Native -36.8282 -73.0347
7982 Gappa 2000 None 9999-01-01 Port Stanley Crypto -51.6921 -57.8589

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