Invasion History
First Non-native North American Tidal Record: 1987First 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
Potentially Misidentified Species
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
SusFedHabitats
General Habitat | Grass Bed | None |
General Habitat | Coarse Woody Debris | None |
General Habitat | Marinas & Docks | None |
General Habitat | Rocky | None |
General Habitat | Vessel Hull | None |
Salinity Range | Polyhaline | 18-30 PSU |
Salinity Range | Euhaline | 30-40 PSU |
Tidal Range | Subtidal | None |
Tidal Range | Low Intertidal | None |
Vertical Habitat | Epibenthic | None |
Tolerances and Life History Parameters
Minimum Temperature (ºC) | -1.8 | Freezing point of seawater, based on range (Saunders and Metaxas 2007) |
Maximum Temperature (ºC) | 26 | Experimental, acclimated at 18 C, temperature increased at 2 C per day (Menon 1972). |
Minimum Salinity (‰) | 27 | Field data, Great Bay NH (Blezard 1999) |
Minimum Reproductive Temperature | 8 | Start of larval settlement, July, Nova Scotia (Saunders and Metaxas 2007) |
Maximum Reproductive Temperature | 21 | Pacific populations, La Jolla CA (conspecificity uncertain, Yoshioka 1982) |
Minimum Duration | 28 | Estimated larval duration, field, Pacific (Schwaniger 1999) |
Maximum Duration | 60 | Estimated larval duration, field, Pacific (Schwaniger 1999) |
Broad Temperature Range | None | Cold temperate-Warm temperate |
Broad Salinity Range | None | Polyhaline-Euhaline |
General Impacts
Economic ImpactsFisheries- 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-ET2 | Bay of Fundy to Cape Cod | Ecological Impact | Competition | ||
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-ET2 | Bay of Fundy to Cape Cod | Ecological Impact | Habitat 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-ET2 | Bay of Fundy to Cape Cod | Ecological Impact | Food/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-ET1 | Gulf of St. Lawrence to Bay of Fundy | Ecological Impact | Competition | ||
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-ET1 | Gulf of St. Lawrence to Bay of Fundy | Ecological Impact | Habitat 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 Impact | Competition | ||
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 Impact | Habitat 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 Impact | Competition | ||
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 Impact | Habitat 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 Impact | Food/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).) | |||||
N100 | Casco Bay | Ecological Impact | Food/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). | |||||
N180 | Cape Cod Bay | Ecological Impact | Competition | ||
Extensive fouling by M. membranacea led to widespread death of kelp (Saccharina and Laminaria spp.) in the Cape Cod Canal (Lambert et al. 1992). | |||||
N180 | Cape Cod Bay | Ecological Impact | Habitat 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-S3 | None | Economic Impact | Fisheries | ||
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-ET1 | Gulf of St. Lawrence to Bay of Fundy | Ecological Impact | Food/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). | |||||
N130 | Great Bay | Ecological Impact | Food/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). | |||||
NH | New Hampshire | Ecological Impact | Competition | ||
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). | |||||
NH | New Hampshire | Ecological Impact | Habitat 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). | |||||
MA | Massachusetts | Ecological Impact | Competition | ||
Extensive fouling by M. membranacea led to widespread death of kelp (Saccharina and Laminaria spp.) in the Cape Cod Canal (Lambert et al. 1992). | |||||
MA | Massachusetts | Ecological Impact | Habitat 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). | |||||
ME | Maine | Ecological Impact | Competition | ||
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). | |||||
ME | Maine | Ecological Impact | Food/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; 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).) | |||||
ME | Maine | Ecological Impact | Habitat 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). |
Regional Distribution Map
Bioregion | Region Name | Year | Invasion Status | Population Status |
---|---|---|---|---|
NEA-II | None | 0 | Native | Established |
NEA-III | None | 0 | Native | Established |
NEA-IV | None | 0 | Native | Established |
AR-V | None | 0 | Native | Established |
NEA-V | None | 0 | Native | Established |
NA-ET2 | Bay of Fundy to Cape Cod | 1987 | Non-native | Established |
NA-ET1 | Gulf of St. Lawrence to Bay of Fundy | 1992 | Non-native | Established |
B-II | None | 0 | Native | Established |
B-I | None | 0 | Native | Established |
SA-I | None | 0 | Crypogenic | Established |
AUS-VIII | None | 1879 | Native | Established |
AUS-VII | None | 0 | Native | Established |
AUS-X | None | 0 | Native | Established |
AR-III | None | 0 | Native | Established |
SEP-A' | None | 0 | Native | Established |
NA-ET3 | Cape Cod to Cape Hatteras | 1990 | Non-native | Established |
N130 | Great Bay | 1989 | Non-native | Established |
M010 | Buzzards Bay | 2000 | Non-native | Established |
M040 | Long Island Sound | 1993 | Non-native | Established |
M020 | Narragansett Bay | 2000 | Non-native | Established |
SEP-B | None | 0 | Native | Established |
NZ-IV | None | 0 | Native | Established |
M026 | _CDA_M026 (Pawcatuck-Wood) | 1990 | Non-native | Established |
N050 | Penobscot Bay | 1998 | Non-native | Established |
N070 | Damariscotta River | 1992 | Non-native | Established |
N100 | Casco Bay | 2004 | Non-native | Established |
N110 | Saco Bay | 1998 | Non-native | Established |
N125 | _CDA_N125 (Piscataqua-Salmon Falls) | 1987 | Non-native | Established |
N135 | _CDA_N135 (Piscataqua-Salmon Falls) | 1987 | Non-native | Established |
N170 | Massachusetts Bay | 2000 | Non-native | Established |
N180 | Cape Cod Bay | 1992 | Non-native | Established |
N140 | Hampton Harbor | 2007 | Non-native | Established |
N120 | Wells Bay | 2007 | Non-native | Established |
N080 | Sheepscot Bay | 2007 | Non-native | Established |
NA-S3 | None | 2002 | Non-native | Established |
NA-S2 | None | 2009 | Non-native | Established |
B-IV | None | 0 | Native | Established |
WA-I | None | 1909 | Crypogenic | Unknown |
Occurrence Map
OCC_ID | Author | Year | Date | Locality | Status | Latitude | Longitude |
---|---|---|---|---|---|---|---|
7902 | None | None | 9999-01-01 | None | Non-native | 41.1259 | -73.2396 |
7903 | USGS Nonindigenous Aquatic Species Program | 1990 | 1990-01-01 | Mohegan Bluffs, Block Island | Non-native | 41.1518 | -71.5567 |
7904 | USGS Nonindigenous Aquatic Species Program 2008 | 1990 | 1990-01-01 | Quonochontaug Beach | Non-native | 41.3318 | -71.7217 |
7905 | MacIntyre et al. 2010) | 2010 | 2010-01-01 | Fort Adams State Park, Newport | Non-native | 41.4782 | -71.3356 |
7906 | MIT Sea Grant 2003 | 2000 | 2000-01-01 | Newport Shipyard, Newport | Non-native | 41.4826 | -71.3256 |
7907 | MIT Sea Grant 2003 | 2000 | 2000-01-01 | Tripp Marina | Non-native | 41.5168 | -71.0856 |
7908 | MIT Sea Grant 2003 | 2000 | 2000-01-01 | Massachusetts Maritime Academy, Bourne | Non-native | 41.7395 | -70.6239 |
7909 | MIT Sea Grant 2003 | 2000 | 2000-01-01 | Sandwich | Non-native | 41.7704 | -70.5036 |
7910 | Whitlach and Osman 2000 | 1998 | 1998-01-01 | Plymouth Harbor | Non-native | 41.9751 | -70.6661 |
7911 | MIT Sea Grant 2003 | 2000 | 2000-01-01 | Quincy | Non-native | 42.2395 | -70.9764 |
7912 | MIT Sea Grant 2003) | 2000 | 2000-01-01 | Boston Harbor | Non-native | 42.3834 | -71.0453 |
7913 | Bell et al. 2005 | 2003 | 2003-01-01 | Boston Harbor Islands | Non-native | 42.3301 | -70.9284 |
7914 | MIT Sea Grant 2003) | 2000 | 2000-01-01 | Salem Harbor | Non-native | 42.5220 | -70.8823 |
7915 | MIT Sea Grant 2003 | 2000 | 2000-01-01 | Gloucester Harbor | Non-native | 42.5959 | -70.6689 |
7916 | MIT Sea Grant 2009 | 2007 | 2007-01-01 | Hampton River Marina, Hampton | Non-native | 42.8995 | -70.8209 |
7917 | Berman et al. 1993; Harris and Tyrell 2000 | 1987 | 1987-01-01 | Isles of Shoals | Non-native | 42.9865 | -70.6120 |
7919 | USGS Nonindigenous Aquatic Species Program 2008 | 1989 | 1989-01-01 | near Fox Pt. | Non-native | 43.1212 | -70.8589 |
7920 | Ruiz et al. unpublished data | 2001 | 2001-01-01 | Badgers Island Marina, Portsmouth | Non-native | 43.0823 | -70.7517 |
7921 | Whitlatch and Osman 2000 | 1998 | 1998-01-01 | York Harbor | Non-native | 43.1306 | -70.6312 |
7922 | Berman et al. 1992; Lambert et al. 1992 | 1987 | 1987-01-01 | Cape Neddick | Non-native | 43.1698 | -70.6034 |
7923 | Whitlatch and Osman 2000 | 1998 | 1998-01-01 | Cape Porpoise | Non-native | 43.3731 | -70.4295 |
7924 | Pratt and Grason 2007 | 2004 | 2004-01-01 | Bailey Island | Non-native | 43.7376 | -69.9937 |
7925 | MIT Sea Grant 2008 | 2007 | 2007-01-01 | Port Harbor Marine, South Portland | Non-native | 43.6415 | -70.2409 |
7926 | MIT Sea Grant 2008 | 2007 | 2007-01-01 | Brewer South Freeport Marina | Non-native | 43.8198 | -70.1095 |
7927 | MIT Sea Grant 2008 | 2007 | 2007-01-01 | Maine DMR Docks, Boothbay Harbor | Non-native | 43.8465 | -69.6348 |
7928 | Lambert et al. 1992; MIT Sea Grant 2008 | 1992 | 1992-01-01 | Darling Maine Center Dock, Wapole | Non-native | 43.9401 | -69.5737 |
7929 | Whitlatch and Osman 2000 | 1998 | 1998-01-01 | Castine Harbor | Non-native | 44.3829 | -68.7989 |
7930 | Whitlatch and Osman 2000 | 1998 | 1998-01-01 | Belfast Bay | Non-native | 44.4281 | -69.0020 |
7931 | MIT Sea Grant 2009 | 2007 | 2007-01-01 | Journey's End Marina, Rockland | Non-native | 44.1045 | -69.1017 |
7932 | Wantanabe et al. 2010 | 2007 | 2007-01-01 | Cape Sable | Non-native | 43.4070 | -65.5050 |
7933 | Watanabe et al. 2009 | 2007 | 9999-01-01 | Sydney | Non-native | 46.1530 | -60.1170 |
7934 | Saunders and Metaxas 2007 | 1992 | 1992-01-01 | St. Margarets Bay | Non-native | 44.7730 | -63.9530 |
7935 | Saunders and Metaxas 2007 | 1992 | 1992-01-01 | Mahone Bay | Non-native | 44.4540 | -64.3530 |
7936 | Watanabe et al. 2009 | 2007 | 2007-01-01 | Canso | Non-native | 45.3660 | -60.9900 |
7937 | Watanabe et al. 2009 | 2007 | 2007-01-01 | None | Non-native | 45.0240 | -62.2130 |
7938 | Fisheries and Oceans Canada 2011 | 2010 | 2010-01-01 | St. Marys Bay | Non-native | 46.9290 | -53.6790 |
7939 | Fisheries and Oceans Canada 2011 | 2009 | 2009-01-01 | Placentia | Non-native | 47.2430 | -53.9270 |
7940 | Fisheries and Oceans Canada 2011 | 2010 | 2010-01-01 | Bay l'Argent | Non-native | 47.5503 | -54.8714 |
7941 | Fisheries and Oceans Canada 2011 | 2010 | 2010-01-01 | Harbour Breton | Non-native | 47.4670 | -55.8440 |
7942 | Gendron et al. 2010) | 2006 | 2006-01-01 | Paspebiac/ | Non-native | 48.0061 | -65.2833 |
7943 | Fisheries and Oceans Canada 2011 | 2002 | 2002-01-01 | Mouse Island | Non-native | 47.6070 | -59.1230 |
7944 | Fisheries and Oceans Canada 2011 | 2002 | 2002-01-01 | None | Non-native | 48.7370 | -58.9260 |
7945 | Fisheries and Oceans Canada 2011 | 2009 | 2009-01-01 | Lark Harbour | Non-native | 49.1050 | -58.4110 |
7946 | Fisheries and Oceans Canada 2011 | 2009 | 2009-01-01 | Point Riche | Non-native | 50.7070 | -57.3910 |
7947 | Caines and Gagnon 2012 | 2009 | 2009-01-01 | Norris Point | Non-native | 49.5530 | -57.8380 |
7948 | Caines and Gagnon 2012 | 2008 | 2008-01-01 | Daniels Harbour | Non-native | 50.2330 | -57.5890 |
7949 | Fisheries and Oceans Canada 2011 | 2009 | 2009-01-01 | Anchor Point | Non-native | 51.3320 | -56.7280 |
7950 | Caines and Gagnon 2012 | 2008 | 2008-01-01 | Green Island Cove | Non-native | 51.3790 | -56.5750 |
7951 | Caines and Gagnon 2012 | 2008 | 2008-01-01 | Red Bay | Non-native | 51.8050 | -56.4000 |
7953 | Fisheries and Ocean Canada 2011 | 2009 | 2009-01-01 | Witless Bay | Non-native | 47.3560 | -52.7460 |
7954 | Fisheries and Ocean Canada 2011 | 2009 | 2009-01-01 | St. Johns | Non-native | 47.5675 | -52.7072 |
7955 | Fisheries and Ocean Canada 2011) | 2009 | 2009-01-01 | Holyrood | Non-native | 47.4340 | -53.1140 |
7956 | Fisheries and Ocean Canada 2011 | 2010 | 2010-01-01 | Dover | Non-native | 48.9900 | -53.9230 |
7957 | Fisheries and Oceans Canada 2011 | 2009 | 2009-01-01 | Lewisporte | Non-native | 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 | Crypogenic | -51.6921 | -57.8589 |
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