Invasion History

First Non-native North American Tidal Record: 1973
First Non-native West Coast Tidal Record: 1973
First Non-native East/Gulf Coast Tidal Record: 1998

General Invasion History:

Caprella mutica is native to the Northwest Pacific. It was first described from Peter the Great Bay, Russia, by Schurin in 1939, and was subsequently found from the Kurile Islands and Akkeshi Bay in Hokkaido Japan (Ashton 2006) south to the Bohai Sea (Laoning Province) and Jiazhou Bay (Shandong Province) in China (Huang 2001). This caprellid has been introduced to the East (Delaware-Newfoundland) and West coasts (California-Alaska) of North America, Europe (from Spain to Norway and Germany), and New Zealand. Caprellids are capable of long-distance dispersal on floating seaweeds or other objects, but ballast water, ship fouling, and the culture of Pacific Oysters (Crassostrea gigas) are likely vectors for the transport of C. mutica to different regions of the world. Many of the occurrence records are associated with aquaculture facilities and other man-made structures, such as breakwaters, marinas, and oil platforms (Platvoet et al. 1995; Willis et al. 2004; Ashton 2006; Page et al. 2006; Ashton et al. 2007; Cook et al. 2007).

North American Invasion History:

Invasion History on the West Coast:

In 1973, Caprella mutica (initially reported as C. acanthogaster) was collected at Field's Landing, in Humboldt Bay, California (Martin 1977, cited by Carlton 1979, Marelli 1981, Boyd et al. 2002). In 1977, it was collected near Oakland in San Francisco Bay (Marelli 1981) and now ranges throughout the South and Central Bay (Cohen et al. 2005). It has been collected at many locations in California, including Elkhorn Slough (in 1978, Marelli 1981), San Diego Bay (in 2001, Fairey et al. 2002, reported as C. acanthogaster), Morro Bay, Channel Islands Harbor, Port Hueneme, Los Angeles-Long Beach Harbors, San Diego Bay (Cohen et al. 2002; Fairey et al. 2002), and oil platforms off Santa Barbara (Page et al. 2006). To the north, it was first collected in Coos Bay, Oregon in 1983 (Carlton 1989; Wonham et al. 2005), Puget Sound, Washington in 1998 and Victoria, British Columbia in 1995 (Cohen et al. 1998; Cohen et al. 2001; Frey et al. 2009). Later it was found to range much farther north, including the Queen Charlotte Islands and Prince Rupert, British Columbia (Frey et al. 2009), and Ketchikan to Kachemak Bay, and Dutch Harbor, in the Aleutian Islands. The timing of this northern invasion is not clear- the earliest Alaskan record is from Sitka in 2001 (Ashton et al. 2008). Pre-existing populations or natural range extensions are conceivable, but extensive collections of caprellids were made in Alaska in the 19th century (US National Museum of Natural History 2012), and in British Columbia in the 20th century (Frey et al. 2009). Potential vectors for C. mutica's transport to the West coast include transplants of Pacific Oysters (Crassostrea gigas), hull fouling, and ballast water.

Invasion History on the East Coast:

The earliest collection of Caprella mutica on the East Coast of North America was in 1998 in Brundenel, on Prince Edward Island, Canada, in the Gulf of St. Lawrence (Locke et al. 2007; Turcotte and Sainte Marie 2009). In 2000, in a survey of southern New England harbors, Caprella mutica was collected in many locations from Gloucester, Massachusetts, south to Newport, Rhode Island. A subsequent survey in 2003 extended the range north to Freeport, Maine, and south to Mystic, Connecticut (MIT Sea Grant 2003). In 2013, established populations of C. mutica were found in the Indian River Inlet, Delaware and Ocean City Inlet, Maryland (Macarena Ros, personal communication, 2013). This caprellid was also collected in Passamaquoddy Bay, New Brunswick, Canada (in 2002, Ashton 2006), Placentia Bay on the south shore of Newfoundland (in 2010, Fisheries and Oceans Canada 2011), and at several more locations in the Gulf of St. Lawrence, including Chaleur Bay (in 2003) and the Magdalen Islands, Quebec (in 2005, Turcotte and Sainte Marie 2009).

Invasion History Elsewhere in the World:

Caprella mutica was first reported in European waters from a surge barrier in Zeeland, and Burghsluis, Netherlands, on the Eastern Scheldt estuary, where it was named as a new species C. macho (Platvoet 1995). In 1999, it was found near a salmon farm in Oban, on the west coast of Scotland (Willis et al. 2004), the Shetland Islands and in Austevoll, Norway (Skifterik 2001; Ashton 2006; Cook et al. 2007). This caprellid is now known from much of the coast of Northwestern Europe, from Le Havre, France (on the English Channel) to Helgoland, Germany (on the North Sea), including the Channel Coast of England, the east and west coasts of Scotland, and the west and east coasts of Ireland (Tierney et al. 2004; Buschbaum and Gutow 2005; Ashton 2006; Arenas et al. 2006; Ashton et al. 2007; Cook et al. 2007; Schückel et al. 2010).

In the Southwest Pacific, Caprella mutica is established near Timaru and Lyttleton on the South Island of New Zealand (Willis et al. 2009).


Description

Caprellid amphipods have a greatly modified body form, when compared to more familiar gammarid amphipods. The body is elongated (giving rise to the name 'skeleton shrimp'), though the abdomen is compressed. The head is partly fused with the first thoracic segment (called Pereonite 1 in amphipods). The head bears a pair of long antennae 1, somewhat shorter antennae. The 1st antennae (A1) have a 3-segmented peduncle, tipped by a flagellum with multiple segments. The 2nd antennae (A2) may be fringed with long setae, and have 3-4 segments in the peduncle, and a shorter flagellum, usually of 2 segments. A mandibular palp of several segments is present in some genera, arising between the antennae, but this is absent in Caprella. There is a small pair of gnathopods, with small grasping claws, with a movable finger (Gnathopod 1) on Pereonite 1. Pereonite 2 bears a much larger pair of gnathopods (Gnathopod 2), which may have conspicuous spines or setae. Pereonites 3 and 4 usually have round or club-shaped gills, while in most species, including Caprella, pereiopods are absent. Pereopods 5, 6, and 7 are roughly equal and hook-like, for climbing and attachment, with 6 segments. Females develop oostegites, plates which form a brood pouch. Males are usually larger than females of the same species. Females and immature males can be hard to identify to species level. (Description from: Barnes 1983; Watling and Carlton, in Carlton 2007).  

Caprella mutica males can grow up to 35 mm, while females grow up to 15 mm (MarLin 2006). Paired dorsal and lateral spines on pereonites 3 to 7 increase in number and size with maturity. Mature males can often be distinguished from other caprellids by naked eye. Immature specimens may only have small paired dorsal spines on pereonite 5. Live specimens of both sexes are bright orange to red in color, with the brood pouch of the female being pale white with dark red dots (Platvoet et al. 1995). There are a number of morphological differences between male and females. Females have no setation on the first and second pereopods, but do have dorsal and lateral spines on pereonites 3 to 7. The pereonites and Gnathopod 2 of females are greatly shortened compared with those of the males. Mature females are distinguished by the developing oostegites and brood pouch. (See further descriptions from: Platvoet et al. 1995; Ashton 2006; MarLin 2006; and Watling and Carlton, in Carlton 2007)


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Arthropoda
Subphylum:   Crustacea
Class:   Malacostraca
Subclass:   Eumalacostraca
Superorder:   Peracarida
Order:   Amphipoda
Suborder:   Caprellidea
Infraorder:   Caprellida
Superfamily:   Caprelloidea
Family:   Caprellidae
Genus:   Caprella
Species:   mutica

Synonyms

Caprella acanthogaster humboldtiensis (Martin, 1977)
Caprella macho (Platvoet, De Bruyne and Gmelig-Meyling, 1995)

Potentially Misidentified Species

Caprella acanthogaster
Carlton (1979) and Marelli (1981) mistakenly applied this name to Caprella mutica in California. Caprella acanthogaster is also native to the northwest Pacific, and has been introduced to Tasmania (Guerra-Garcia and Takeuchi 2004), but is not known from North American or European waters.

Caprella alaskana
NE Pacific native, California to Alaska

Caprella kennerlyi
NE Pacific native, California to Alaska

Caprella laeviuscula
NE Pacific native, California to Alaska

Caprella linearis
N Atlantic native, New England-Labrador, Arctic Russia-Spain, Alaska

Ecology

General:

Life History – The males are large and slender, armed with larger gnathopods, probably an adaptation to compete for females and to guard themselves during molting. The embryos are brooded by the female in an egg-pouch formed by large plates (oostegites) on the 3rd and 4th pereionites (Turcotte and Sainte Marie 2009). Development is direct, with the newborn juveniles having the general form of adults. Females reach maturity at Instar VII (the 6th molt), an average of 53 days after birth (cultured at 14⁰C). The average lifespan was 90-120 days with most females producing two broods before death (Cook et al. 2007). However, estimated brooding time and lifespan varies greatly in the field, with varying temperature and food conditions (Turcotte and Sainte Marie 2009). The importance of maternal care in C. mutica is unclear (Boos et al. 2011). In culture, newborn juvenile C. mutica may cling to the females after birth, and remain nearby for several days, but disperse within a week. Brooding females and females with newborns are more likely to fight with males than non-reproducing females (Matthews 2008). The population cycle is strongly seasonal in the Sea of Japan and in Chaleur Bay, Quebec, with peaks of reproduction in spring (April-May) and summer (June-August), and two generations per year (Turcotte and Sainte Marie 2009). In Scotland, with somewhat higher winter temperatures, (minimum of 7⁰C) reproduction occurred year-round, but populations peaked in July-August (Ashton 2006). In culture, C. mutica was able to reproduce at 4-26⁰C, but reproduction was optimal at 16⁰C and impaired at the upper and lower limits (Boos et al. 2011).

Ecology – Caprellids can feed in a variety of ways, including filtering small particles from the water, browsing on small filamentous algae, scraping tissue from large algae, scavenging, and predation (Turcotte and Sainte Marie 2009). Caprella mutica appears to be capable of using all these modes of feeding, which may contribute to its success as an invader (Cook et al. 2007, Turcotte and Sainte Marie 2009; Cook et al. 2010; Best et al. 2013). The high abundance of C. mutica found around salmon farms may be due to direct consumption of salmon feed, as well as feeding on enhanced densities of phytoplankton stimulated by nutrients from the aquaculture operation (Boos et al. 2011). It has been found clinging to vegetation, hydroids, bryozoans, and manmade structures (ropes, nets, etc.) (Ashton 2006; Willis et al. 2004; Maciejeski 2008). It tolerates wide ranges of temperature, 2 - 25⁰C, and salinities as low as 11 PSU in the field in the northern Sea of Japan (Schevchenko et al. 2004, cited by Turcotte and Sainte Marie 2009). In laboratory experiments, the temperature range was similar, 2-26⁰C, but the lower salinity limit was higher at 15 PSU. It showed little mortality at the highest salinity tested, 40 PSU (Ashton et al. 2007). In North Sea, off Netherlands and Belgium, C. mutica was dominant on intertidal and floating artificial substrates, in waters with high densities of suspended particulate matter, and less than 17 m depth, while the native C.linearis dominated on fixed, subtidal substrates in deeper water (Coolen et al. 2016). Caprella mutica appears to have sufficient tolerance and flexibility of habitat, feeding, and life history to colonize much of the world's temperate waters (Ashton 2006; Boos et al. 2011). 

Food:

Phytoplankton, seaweed, detritus

Consumers:

Fishes

Competitors:

Caprella californica, Caprella linearis

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatVessel HullNone
General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-2Based on field distribution, laboratory animals tolerated 2 C, lowest tested (Ashton 2006)
Maximum Temperature (ºC)28Experimental, 48 h LT 50 (Ashton 2006)
Minimum Salinity (‰)14.6Experimental, 48 h LC 50, reduced activity below 18 ppt (Ashton 2006)
Maximum Salinity (‰)40Experimental, highest tested (Ashton 2006)
Minimum Reproductive Temperature4Lowest tested (Ashton 2006). Hatchlings maintained at 4 C died after 4 months (Boos et al. 2011), but winter hatchlings probably would survive a normaly seasonal cycle with spring warming.
Maximum Reproductive Temperature20Highest tested (Boos et al. 2011)
Maximum Length (mm)50Nihsimura 1995, males, in Japan, cited by Cook et al. 2007
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

In its invaded range, Caprella mutica has achieved extraordinary densities, especially on man-made structures. Studies of its economic and ecological impacts are limited, but observations indicate that C. mutica can affect aquaculture operations, displace native caprellids, and affect the feeding of native fishes (Ashton 2006; Page et al. 2007; Shucksmith et al. 2009; Turcotte and Sainte Marie 2009).

In Price Edward Island, Canada, high densities of caprellids (C. mutica and C. linearis) were shown to inhibit the settlement of the tunicate Ciona intestinalis (Collin and Johnson 2014). Since the tunicate appears to be locally invasive, and interfering with mussel aquaculture, there may be a potential for biocontrol by encouraging caprellid populations (Collin and Johnson 2014).

Regional Impacts

NEP-VIPt. Conception to Southern Baja CaliforniaEcological ImpactFood/Prey
On oil platforms off Santa Barbara, the caprellid community was dominated by Caprella mutica, which was a major component of the diet of a native fish, the Painted Greenling (Oxylebius pictus), which was very abundant at the platform (Page et al. 2007).
P065_CDA_P065 (Santa Barbara Channel)Ecological ImpactFood/Prey
On oil platforms off Santa Barbara, the caprellid community was dominated by Caprella mutica, which was a major component of the diet of a native fish, the Painted Greenling (Oxylebius pictus), which was very abundant at the platform (Page et al. 2007).
NEA-IIINoneEcological ImpactCompetition
In laboratory experiments in Scotland, C. mutica displaced the native caprellids C. linearis and Pseudoprotella phasma from natural (hydroid) and artificial habitat (plastic mesh) patches on fouling plates. Displacement of C. linearis occurred even when the density of the native was 10X that of C. mutica (Shucksmith et al. 2009).
NEA-IIINoneEconomic ImpactFisheries
Mussel farmers observed reduced settlement of spat during periods where C. mutica was most abundant; however a causal connection could not be confirmed (Ashton 2006).
NA-S3NoneEconomic ImpactFisheries
Field and laboratory work (unpublished) indicates that high densities of C. mutica interfere with settlement of mussel spat (Turcotte and Sainte Marie 2009).
NA-S3NoneEcological ImpactCompetition
On fouling plates, high densities of Caprella mutica and the cryptogenic C. linearis inhibit the settlement of the tunicate Ciona intestinalis, which is locally invasive in the Gulf of St. Lawrence. One possible mechanism is physical disturbance or chemical avoidance of caprellids by settling tunicate larvae (Collin and Johnson 2014).
NA-S3NoneEcological ImpactPredation
A negative correlation between newly settled tunicates (Ciona intestinalis) and caprellids (C. mutica and C. linearis) on fouling plates in Prince Edward Island estuaries may be the result of predation by caprellids on the larvae. While the two caprellids were equally abundant, C. mutica's larger size suggests that it made the largest contribution to the interaction (Collin and Johnson 2014).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactPredation
In Bodega Harbor, caging experiments and feeding trials showed that Caprella mutica was a significant predator on recruits of Ciona intestinalis (Rius et al. 2014).
P112_CDA_P112 (Bodega Bay)Ecological ImpactPredation
In Bodega Harbor, caging experiments and feeding trials showed that Caprella mutica was a significant predator on recruits of Ciona intestinalis (Rius et al. 2014).
P090San Francisco BayEcological ImpactCompetition
Caprella mutica was less tolerant of low salinity (LC50 of 17-21 PSU) than C. californica (LC50 of 16 PSU), and died out more quickly after low-salinity events in San Francisco Bay, but recolonized more rapidly due to faster maturity and higher fecundity (Desmet 2011).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NWP-4a None 0 Native Estab
NEP-V Northern California to Mid Channel Islands 1977 Def Estab
NEP-IV Puget Sound to Northern California 1973 Def Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1995 Def Estab
NA-ET2 Bay of Fundy to Cape Cod 2000 Def Estab
NEA-II None 1995 Def Estab
AR-V None 1999 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1998 Def Estab
NWP-4b None 0 Native Estab
NEP-VI Pt. Conception to Southern Baja California 2000 Def Estab
NEA-III None 2000 Def Estab
P130 Humboldt Bay 1973 Def Estab
M040 Long Island Sound 2003 Def Estab
M010 Buzzards Bay 2000 Def Estab
P050 San Pedro Bay 2000 Def Estab
P170 Coos Bay 1983 Def Estab
M020 Narragansett Bay 2000 Def Estab
P062 _CDA_P062 (Calleguas) 2000 Def Estab
P020 San Diego Bay 2001 Def Estab
P070 Morro Bay 2001 Def Estab
P080 Monterey Bay 1978 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 2002 Def Estab
P090 San Francisco Bay 1977 Def Estab
P110 Tomales Bay 2001 Def Estab
P112 _CDA_P112 (Bodega Bay) 2001 Def Estab
P290 Puget Sound 1998 Def Estab
N100 Casco Bay 2003 Def Estab
N170 Massachusetts Bay 2000 Def Estab
N180 Cape Cod Bay 2000 Def Estab
NWP-5 None 0 Native Estab
NA-S3 None 1998 Def Estab
NZ-IV None 2004 Def Estab
NEP-II Alaska south of Aluetians to the Alaskan panhandle 2002 Def Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 2007 Def Estab
N130 Great Bay 2007 Def Estab
N140 Hampton Harbor 2007 Def Estab
N070 Damariscotta River 2007 Def Estab
N050 Penobscot Bay 2007 Def Estab
N080 Sheepscot Bay 2007 Def Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 2010 Def Estab
N120 Wells Bay 2011 Def Estab
N125 _CDA_N125 (Piscataqua-Salmon Falls) 2011 Def Estab
B-I None 2011 Def Estab
M026 _CDA_M026 (Pawcatuck-Wood) 2010 Def Estab
N195 _CDA_N195 (Cape Cod) 2000 Def Estab
M100 Delaware Inland Bays 2013 Def Estab
M120 Chincoteague Bay 2013 Def Estab
NEA-V None 2013 Def Estab
P030 Mission Bay 2011 Def Estab
WA-IV None 2015 Def Estab
NEA-IV None 2018 Def Estab
M060 Hudson River/Raritan Bay 2019 Def Estab
P040 Newport Bay 2017 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
7400 MIT Sea Grant 2003 2003 2003-01-01 Brewers Yacht Yard, Mystic Def 41.3543 -71.9665
7401 MacIntyre et al. 2010 2010 2010-07-25 Pt. Judith Marina Def 41.3788 -71.5169
7403 MIT Sea Grant 2003 2000 2000-01-01 Coasters Harbor Island Def 41.5107 -71.3270
7404 MIT Sea Grant 2003 2000 2000-01-01 Fort Getty Pier, Jamestown Def 41.4915 -71.3983
7405 MIT Sea Grant 2003 2000 2000-01-01 Wickford Marina, Wickford Def 41.5754 -71.4423
7406 MIT Sea Grant 2003 2000 2000-01-01 Colt State Park, Bristol Def 41.6698 -71.3009
7407 MIT Sea Grant 2003 2000 2000-01-01 Massachusetts Maritime Academy, Bourne Def 41.7395 -70.6239
7408 MIT Sea Grant 2003 2000 2000-01-01 New Bedford State Pier Def 41.5932 -70.8870
7409 MIT Sea Grant 2003 2000 2000-01-01 Tripps Marina, Def 41.5168 -71.0856
7410 MIT Sea Grant 2003 2000 2001-01-01 Woods Hole Coast Guard Station Def 41.5196 -70.6661
7411 MIT Sea Grant 2003 2000 2000-01-01 Barnstable Harbor Def 41.7168 -70.2661
7412 MacIntyre et al. 2010 2010 2010-01-01 Sandwich Marina Def 41.7704 -70.5036
7413 MIT Sea Grant 2003 2000 2000-01-01 Duxbury Harbor, Cape Cod Bay Def 41.9876 -70.6492
7414 MIT Sea Grant 2003 2000 2000-01-01 Bay Pointe Marina, Quincy Def 42.2918 -70.9745
7415 MIT Sea Grant 2003 2000 2000-01-01 Rowes Wharf, Boston Def 42.3570 -71.0409
7416 MIT Sea Grant 2003 2000 2000-01-01 Hawthorne Cove Marina Def 42.5220 -70.8823
7417 MIT Sea Grant 2003 2000 2000-01-01 Cape Ann Marina, Gloucester Def 42.5959 -70.6689
7418 MIT Sea Grant 2009 2007 2007-01-01 Hampton River Marina Def 42.8995 -70.8209
7419 Harris and Dijkstra 2007 2007 2007-01-01 Isles of Shoals Def 42.9865 -70.6120
7420 MacIntyre et al. 2010 2010 2010-01-01 UNH Coastal Marine Lab, Newcastle Def 43.0723 -70.7162
7421 MIT Sea Grant 2008 2007 2007-01-01 Wentworth by the Sea Marina Def 43.0523 -70.7334
7422 MIT Sea Grant 2012 2011 2011-01-01 York Harbor Def 43.1368 -70.6456
7423 MIT Sea Grant 2012 2011 9999-01-01 Wells Harbor Def 43.3215 -70.5595
7424 MIT Sea Grant 2003 2003 9999-01-01 Brewer Freeport Marina, South Freeport Def 43.8198 -70.1095
7425 MIT Sea Grant 2009 2007 2007-01-01 MDMR docks, Boothbay Harbor Def 43.8465 -69.6348
7426 MIT Sea Grant 2009) 2007 2007-01-01 Darling Marine Center Dock Def 43.9401 -69.5737
7427 MIT Sea Grant 2009 2007 2007-01-01 Journey's End Marina, Rockland Def 44.1045 -69.1017
7428 Ashton 2006 2000 2000-01-01 Passamaquoddy Bay Def 45.0833 -67.0833
7429 Fisheries and Oceans Canada 2011 2010 2010-01-01 Bay L'Argent Def 47.5458 -54.8828
7430 Fisheries and Oceans Canada 2011 2010 2010-01-01 Barren Island Def 47.7150 -54.0840
7431 Turcotte and Sainte Marie 2009 2004 2004-01-01 Caribou Def 45.8500 -62.6290
7433 Turcotte and Sainte Marie 2009 2004 2004-01-01 Magdalen Islands Def 47.4483 -61.7522
7434 Ashton 2006 2004 2004-01-01 Cascapedia Def 48.2860 -65.8680
7435 Locke et al. 2007 1998 1998-01-01 Brudenell Def 46.1831 -62.4820
7437 Cohen et al. 2002 2000 2000-01-01 Pilot's Dock, Pier F, Long Beach Def 33.7472 -118.2156
7445 Cohen et al. 2005 2004 2004-05-26 Richmond Marina Def 37.9139 -122.3542
7452 Carlton 1979; Marelli 1981 1973 1973-01-01 Field's Landing Def 40.7246 -124.2151
7453 Carlton 1989, Wonham et al. 2005 1983 1983-01-01 Coos Bay Def 43.3406 -124.3219
7454 Cohen et al. 2001 2000 2000-05-20 Steamboat Island, Totten Inlet Def 47.1850 -122.9395
7455 Frey et al. 2009 1995 1995-01-01 Victoria Def 48.4222 -123.3657
7456 Frey et al. 2009 2008 2008-01-01 Barkley Sound Def 48.9000 -125.0800
7457 Frey et al. 2009 2006 2006-09-01 Nanaimo Def 49.2100 -123.9600
7458 Frey et al. 2009 2006 2006-09-01 Campbell River Def 50.0300 -125.2400
7459 Frey et al. 2009 2007 2007-10-01 Broughton Archipelago Def 50.7900 -126.6900
7460 Frey et al. 2009 2007 2007-09-01 Cumshewa Inlet Def 53.0200 -131.9100
7461 Frey et al. 2009 2007 2007-09-01 Hecate Strait Def 54.0100 -132.1400
7462 Frey et al. 2009 2007 2007-10-01 Prince Rupert Def 54.2900 -130.3500
7463 Ashton et al. 2008 2003 2003-01-01 Ketchikan Harbor Def 55.3900 -131.7400
7464 Ashton et al. 2008 2001 2001-01-01 Thomsen Harbor, Sitka Def 57.0500 -135.3500
7465 Ashton et al. 2008 2008 2008-01-01 City Spit Dock, Dutch Harbor, Unalaska Island Def 53.9100 -166.5100
7466 Ashton et al. 2008 2003 2003-01-01 Seldovia Harbor, Kachemak Bay Def 59.6100 -151.4100
7467 Ashton 2006 None 9999-01-01 Kunashir, Kurile Islands Native 44.1167 145.8500
7468 Ashton 2006 None 9999-01-01 Shikotan, Kurile Islands Native 43.8000 146.7500
7469 Ashton 2006 None 9999-01-01 Akkeshi Bay Native 43.0500 144.9000
7470 Ashton 2006 None 9999-01-01 Poss'yet Bay Native 42.5000 130.9167
7471 Huang 2001 None 9999-01-01 Dalian, Liaoning Province Native 38.9208 121.6392
7472 Huang 2001 None 9999-01-01 Qingdao Native 36.0667 120.3833
7473 Ashton 2006 2004 2004-01-01 Timaru Def -44.4000 171.2500
7474 Ashton 2006 2005 2005-01-01 Lyttleton Def -43.6000 172.7200
7475 Willis et al. 2009 2007 2007-01-01 Port Ligar Def -40.9225 173.9538
7476 Willis et al. 2009 2007 2007-01-01 Waihinau Bay Def -40.9500 173.9667
7477 Willis et al. 2004 2000 2000-01-01 Oban Def 56.4515 -5.4555
7478 Ashton et al. 2006 2006 2006-01-01 Craobh Haven Def 56.2150 -5.5567
7479 Tierney et al. 2004 2003 2003-07-16 Bertraghboy Bay Def 53.3883 -9.8528
7480 Breton 2004, cited by Ashton 2006 2004 2004-12-30 Le Havre Def 49.4900 0.1000
7481 Arenas et al. 2006 2004 2004-09-04 Poole Harbor (English Channel) Def 50.6958 -1.9886
7482 Ashton 2006 2006 2006-01-01 Southampton Def 50.8970 -1.4042
7483 Ashton 2006 2002 2002-01-01 Harwich Def 51.9550 1.2625
7484 Ashton et al. 2006 2006 2006-01-01 Port Edgar Def 55.9950 -3.4100
7485 Minchin and Holmes 2006 2005 2005-01-01 Dublin Harbour Def 53.3478 -6.2597
7486 Ashton et al. 2006 2006 2006-01-01 Peterhead Def 57.4900 -1.7867
7487 Kerckhof et al. 2007 1998 1998-01-01 Zeebrugge harbour Def 51.3333 3.3000
7488 Platvoet, 1995 1995 1995-01-01 Burghsluis Def 51.6833 3.7500
7489 Buschbaum and Gutow 2005 2004 2004-01-01 List, Sylt (island) Def 55.0167 8.4333
7490 Buschbaum and Gutow 2005 2004 2004-01-01 Helgoland Def 54.1833 7.8833
7491 Schuckel et al. 2010 2009 2009-01-01 Wilhelmshaven Def 53.5167 8.1333
7492 Boos et al. 2011 2005 2005-01-01 offshore windfarm Def 55.2500 8.0000
7493 Skifterik 2001; Ashton 2006 1999 1999-01-01 Austevoll Def 60.0378 5.2683
7494 Ashton 2006 2002 2002-01-01 Alesund Def 62.4778 6.1903
7495 Ashton 2006 2003 2003-01-01 Shetland Islands Def 60.3038 -1.2689
767322 Ruiz et al., 2015 2012 2012-08-13 Coast Guard, Bodega Bay, California, USA Def 38.3126 -123.0512
767329 Ruiz et al., 2015 2012 2012-08-14 Spud Point South, Bodega Bay, California, USA Def 38.3281 -123.0574
767335 Ruiz et al., 2015 2012 2012-08-14 Spud Point North, Bodega Bay, California, USA Def 38.3301 -123.0572
767345 Ruiz et al., 2015 2012 2012-08-21 Lucas/Tides, Bodega Bay, California, USA Def 38.3284 -123.0445
767352 Ruiz et al., 2015 2012 2012-08-21 Porto Bodega, Bodega Bay, California, USA Def 38.3333 -123.0525
767363 Ruiz et al., 2015 2012 2012-08-22 Tomales-Marshall, Bodega Bay, California, USA Def 38.1514 -122.8888
767374 Ruiz et al., 2015 2012 2012-08-21 Tomales-Nick's Cove, Bodega Bay, California, USA Def 38.1980 -122.9222
767385 Ruiz et al., 2015 2012 2012-08-15 Tomales- Call Box 401, Bodega Bay, California, USA Def 38.1793 -122.9104
767392 Ruiz et al., 2015 2012 2012-08-16 Tomales-SNPS, Bodega Bay, California, USA Def 38.1359 -122.8719
767435 Ruiz et al., 2015 2013 2013-07-23 Marina Village, Mission Bay, CA, California, USA Def 32.7605 -117.2364
767455 Ruiz et al., 2015 2013 2013-07-29 Mission Bay Yacht Club, Mission Bay, CA, California, USA Def 32.7778 -117.2485
767475 Ruiz et al., 2015 2013 2013-08-04 Bahia Resort Marina, Mission Bay, CA, California, USA Def 32.7731 -117.2478
767522 Ruiz et al., 2015 2013 2013-08-03 Mission Bay Sport Center, Mission Bay, CA, California, USA Def 32.7857 -117.2495
767550 Ruiz et al., 2015 2013 2013-08-02 The Dana Marina, Mission Bay, CA, California, USA Def 32.7671 -117.2363
767577 Ruiz et al., 2015 2013 2013-08-30 201 Main, Morro Bay, CA, California, USA Def 35.3564 -120.8474
767590 Ruiz et al., 2015 2013 2013-08-27 City Harbor, Morro Bay, CA, California, USA Def 35.3709 -120.8582
767602 Ruiz et al., 2015 2013 2013-09-05 Launch Ramp, Morro Bay, CA, California, USA Def 35.3577 -120.8508
767611 Ruiz et al., 2015 2013 2013-08-29 Moorings, Morro Bay, CA, California, USA Def 35.3619 -120.8548
767624 Ruiz et al., 2015 2013 2013-08-31 Morro Bay Marina, Morro Bay, CA, California, USA Def 35.3641 -120.8532
767631 Ruiz et al., 2015 2013 2013-08-28 Sealion Dock, Morro Bay, CA, California, USA Def 35.3658 -120.8555
767642 Ruiz et al., 2015 2013 2013-09-03 State Park Marina, Morro Bay, CA, California, USA Def 35.3459 -120.8423
767654 Ruiz et al., 2015 2013 2013-09-04 Tidelands, Morro Bay, CA, California, USA Def 35.3602 -120.8521
767663 Ruiz et al., 2015 2013 2013-07-16 Naval Base Point Loma, San Diego Bay, CA, California, USA Def 32.6886 -117.2343
767758 Ruiz et al., 2015 2013 2013-07-26 Pier 32 Marina, San Diego Bay, CA, California, USA Def 32.6516 -117.1077
767799 Ruiz et al., 2015 2011 2011-09-15 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9117 -122.3494
767819 Ruiz et al., 2015 2011 2011-09-20 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8067 -122.4432
767831 Ruiz et al., 2015 2011 2011-09-14 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5880 -122.3160
767852 Ruiz et al., 2015 2011 2011-09-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6725 -122.3864
767877 Ruiz et al., 2015 2011 2012-09-15 Berkeley Marina, San Francisco Bay, CA, California, USA Def 37.8758 -122.3181
767887 Ruiz et al., 2015 2011 2012-09-19 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
767902 Ruiz et al., 2015 2011 2011-09-21 South Beach Harbor, San Francisco Bay, CA, California, USA Def 37.7797 -122.3871
767914 Ruiz et al., 2015 2011 2011-09-20 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7947 -122.2822
767928 Ruiz et al., 2015 2011 2011-09-22 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
767963 Ruiz et al., 2015 2011 2011-09-12 Corinthian Yacht Club, San Francisco Bay, CA, California, USA Def 37.8103 -122.3228
767977 Ruiz et al., 2015 2012 2012-08-24 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9134 -122.3523
768000 Ruiz et al., 2015 2012 2012-08-23 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8609 -122.4853
768017 Ruiz et al., 2015 2012 2012-08-28 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8071 -122.4341
768032 Ruiz et al., 2015 2012 2012-08-27 Port of San Francisco Pier 31, San Francisco Bay, CA, California, USA Def 37.8078 -122.4060
768047 Ruiz et al., 2015 2012 2012-08-31 Antioch Marina, San Francisco Bay, CA, California, USA Def 38.0203 -121.8211
768054 Ruiz et al., 2015 2012 2012-09-11 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7676 -122.2869
768076 Ruiz et al., 2015 2012 2012-08-30 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6633 -122.3817
768100 Ruiz et al., 2015 2012 2012-08-29 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3174
768123 Ruiz et al., 2015 2012 2012-09-04 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5023 -122.2130
768147 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9736 -122.4802
768165 Ruiz et al., 2015 2012 2012-09-05 Port of Oakland, San Francisco Bay, CA, California, USA Def 37.7987 -122.3228
768188 Ruiz et al., 2015 2012 2012-09-07 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7940 -122.2787
768227 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6962 -122.1919
768244 Ruiz et al., 2015 2012 2012-09-12 Emeryville, San Francisco Bay, CA, California, USA Def 37.8396 -122.3133
768267 Ruiz et al., 2015 2013 2013-08-15 Ballena Isle Marina, San Francisco Bay, CA, California, USA Def 37.7656 -122.2858
768290 Ruiz et al., 2015 2013 2013-08-20 Coyote Point Marina, San Francisco Bay, CA, California, USA Def 37.5877 -122.3163
768310 Ruiz et al., 2015 2013 2013-08-22 Jack London Square Marina, San Francisco Bay, CA, California, USA Def 37.7926 -122.2746
768328 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9723 -122.4829
768349 Ruiz et al., 2015 2013 2013-08-13 Oyster Point Marina, San Francisco Bay, CA, California, USA Def 37.6639 -122.3821
768370 Ruiz et al., 2015 2013 2013-08-14 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5024 -122.2134
768393 Ruiz et al., 2015 2013 2013-08-19 Richmond Marina Bay Yacht Harbor, San Francisco Bay, CA, California, USA Def 37.9138 -122.3522
768413 Ruiz et al., 2015 2013 2013-08-12 San Francisco Marina, San Francisco Bay, CA, California, USA Def 37.8078 -122.4354
768445 Ruiz et al., 2015 2013 2013-08-16 Sausalito Marine Harbor, San Francisco Bay, CA, California, USA Def 37.8611 -122.4851

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