Invasion History

First Non-native North American Tidal Record: 1890
First Non-native West Coast Tidal Record: 1890
First Non-native East/Gulf Coast Tidal Record: 2004

General Invasion History:

Urosalpinx cinerea is native to the Atlantic Coast of North America, from Florida to at least Cape Cod, Massachusetts. It occurs further north in Nova Scotia, the Magdalen Islands (Quebec), and Prince Edward Island, in scattered localized populations (Verrill and Smith 1873; Carriker 1955; Bousfield 1960; Abbott 1974). Oyster transplants from southern waters in the 19th century may have extended the range, and/or increased the abundance of U. cinerea in New England and the Maritime provinces (Carriker 1955). However, shells of this whelk were found in excavations of a 5,000 yr-old fish weir in Boston (Johnson et al. 1942). Urosalpinx cinerea may have extended its range north during a warm post-glacial period, 6000-3000 yr B.P., leaving relict populations as the climate cooled, a pattern seen in many marine and terrestrial organisms in the Maritime provinces (Bousfield and Thomas 1975; Pielou 1991).

The Atlantic Oyster Drill has been introduced to the West Coast of North America, from California to British Columbia. It is also established in England where it is abundant in bays where Eastern Oysters (Crassostrea virginica) were unsuccessfully planted in the 19th century (Carriker 1955; Cole 1942; Carlton 1992). Recently, this drill has appeared in the Netherlands (Faase and Ligthart 2009) and on the Gulf Coast of Florida (Robinson and Dillon 2008).

North American Invasion History:

Invasion History on the West Coast:

On the West Coast, Urosalpinx cinerea was first reported in 1890 from beds of planted Eastern Oysters (Crassostrea virginica) in Belmont, San Francisco Bay, California (CA) (Townsend 1890, cited by Carlton 1979). This snail is abundant and widespread in the central and south portions of the Bay, from Alameda southward (Carriker 1955; Cohen and Carlton 1995; Cohen et al. 2005). Other introductions on the West Coast have been highly localized; in part because this species lacks planktonic larvae, and has limited dispersal ability. On the California coast, U. cinerea was first found in Tomales Bay by 1935 (Bonnot 1935, cited by Carlton 1979), Humboldt Bay in 1941 (Carriker 1955; Carlton 1979), Elkhorn Slough in 1945 (Burch 1945, cited by Carlton 1979), and Newport Bay in 1957 (Carlton 1979). The Elkhorn Slough population is extinct (Carlton 1979; Wasson et al. 2001) and the Humboldt Bay population was reported to 'no longer persist', but this drill was collected there in 2000 by Boyd et al. (2002). The Newport Bay population may have been transported in boat fouling or with seaweed used to pack bait, since this Southern California bay has no history of oyster introduction (Carlton 1979).

Further north, U. cinerea was reported from Puget Sound, Washington (WA) (Galtsoff 1929, cited by Carlton 1979; Carriker 1955; Hanna 1966); Boundary Bay, WA (Sherwood 1931, cited by Carlton 1979); Ladysmith (Vancouver Island), British Columbia (Elsey 1933, cited by Carlton 1979); and Willapa Bay, WA (in 1948, Hanna 1966). Locations in Puget Sound included Oakland Bay, Oyster Bay-Totten Inlet, Mud Bay-Eld Inlet, Nisqually Flats, and Frinton Flats (Carriker 1955). The Ladysmith population is extinct and the status of Puget Sound populations is uncertain (Eissinger 2009). Ruesink (in Boersma et al. 2006) mentions only two established populations in the Pacific Northwest: Willapa Bay, 'thriving' and Boundary Bay, 'dwindling'.

Invasion History on the East Coast:

In the 19th century the depletion of oyster beds in New York and New England led to massive northward transplantation of oysters from New Jersey, Delaware, and Chesapeake Bays (Carlton and Mann 1996). This movement of oysters likely carried a number of other species, but the limited number of biologists and collectors at that time, makes it difficult to document alteration to the ranges of species. Fishermen and naturalists noted the appearance of oyster drills, in new locations or an increase in their abundance, in the Hudson estuary, New York (NY); Great South Bay, NY; Long Island Sound, NY; Narragansett Bay, Rhode Island; Cape Cod, Massachusetts (MA); and Massachusetts Bay, MA. Urosalpinx cinerea was widely believed to have been introduced with the transplanted oysters (Carriker 1955). However, Verrill and Smith (1873) mention a population in Casco Bay, Maine and an 'isolated colony' in the Gulf of St. Lawrence. It is likely that U. cinerea in the Northeast consists of a mixture of native, scattered relict populations and introduced populations from southern bays (Carriker 1955).

Invasion History on the Gulf Coast:

Two species of Urosalpinx are native to the Gulf coast of Florida (FL), U. perrugata (Gulf Oyster Drill) and U. tampensis (Tampa Drill) (Abbott 1974). In 2004, Robinson and Dillon (2008) found a large population of U. cinerea living sympatrically with the two native species, on oyster reefs off Cedar Key, FL. No hybridization was observed. The Atlantic Oyster Drill may have been introduced with transplants of oysters or with aquaculture of Atlantic Hard Clams (Mercenaria mercenaria) (Robinson and Dillon 2008).

Invasion History Elsewhere in the World:

Urosalpinx cinerea was first reported in the Northeast Atlantic in Crouch Estuary, Essex, England in 1927 (Cole 1942; Eno et al. 1997). It is now found in a limited area on the east coast of England, in the Crouch and Blackwater Estuaries, as well as, other estuaries in Essex and Kent (Eno et al. 1997). The snail's abundance greatly increased by the 1950s, reaching densities of 15,000-20,000 per hectare. In the 1980s, populations of U. cinerea declined drastically, bringing populations in several estuaries close to extinction. The cause appears to be tributyltin (TBT) pollution from anti-fouling paint, resulting in imposex malformation of female reproductive organs, developing male characteristics, resulting in sterility (Gibbs et al. 1991). In 2007, breeding populations of the Atlantic Oyster Drill were discovered in the Oosterschelde estuary, Netherlands. The banning of TBT may have facilitated the invasion in the Netherlands, and may permit the recovery of English populations (Gibbs et al. 1991; Faasse and Ligthart 2009).


Description

Urosalpinx cinerea is a mid-sized predatory snail. Its shell is dextrally coiled and has a pointed spire. The siphonal canal is moderately long, open and wide. Each whorl has 9 to 11 rounded axial ribs, prominent at the shoulders of the whorls, and numerous spiral cords. The outer lip is slightly thickened and sometimes bears 2-6 small teeth. Adult shells usually reach 25-35 mm, but some populations, particularly on the Atlantic coast of Virginia, tend to produce larger specimens around 50-61 mm long. The shell is usually yellowish or grayish white, often with irregular brown spiral bands. Eggs are laid in yellowish, vase-shaped capsules. Description from: Carriker 1955, Abbott 1974, Gosner 1978, and McLean 2007.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Gastropoda
Subclass:   Prosobranchia
Order:   Neogastropoda
Family:   Muricidae
Genus:   Urosalpinx
Species:   cinerea

Synonyms

Buccinum plicosum (Gould, 1841)
Fusus cinereus (Say, 1822)
Fusus imbricatus (De Kay, 1843)
Urosalpinx aitkinae (Wheat, 1913)
Urosalpinx follyensis (B. B. Baker, 1951)
Urosalpinx cinereus (Say, 1822)

Potentially Misidentified Species

Ocenebra erinacea
European Sting Winkle

Urosalpinx perrugata
Gulf Oyster Drill, found on the East and Gulf coasts of Florida (Abbott 1974; Rosenberg 2014).

Urosalpinx tampaensis
Tampa Oyster Drill, found on the East and Gulf coasts of Florida (Abbott 1974; Rosenberg 2014).

Ecology

General:

Urosalpinx cinerea, the Atlantic Oyster Drill, has separate sexes. It reaches maturity at about 15 months (Delaware) to 3 years (England) (Cole 1942; Carriker 1955). Fertilization is internal. Eggs are laid on rocks or shells in vase-shaped egg capsules, which contain 10-13 eggs. The number of capsules per spawning ranged from 4 to 271, but 25-100 appears to be a more normal range (Manzi 1970). The eggs hatch in about 22-78 days (Ganaros 1958), into miniature snails, 1.0-1.5 mm long (Carriker 1955; Manzi 1970; Cohen 2005). Development is direct, so dispersal is limited.

Atlantic Oyster Drills inhabit oyster beds and rocky intertidal areas, down to about 8 m depth (Cole 1942; Abbot 1974). In Georgia, they are common in the lower portions of intertidal oyster beds, but rarely reach the upper regions (Walker 1971). They tolerate salinities as low as 15 PSU, but need at least 20 PSU for successful egg development.

Urosalpinx cinerea feeds on barnacles, mussels, oysters, and other bivalves, including oysters and mussels. The drill rasps a circular hole in the prey shell and digests its contents. The size of the prey selected tends to be roughly equal with the shell length of the gastropod (Carriker 1955; Harding et al. 1999). Locally important bivalve prey include mussels (Mytilus spp.), Eastern Oysters (Crassostrea virginia), introduced Pacific Oysters (Crassostrea gigas), native Olympic Oysters (Ostrea lurida), and European Oysters (Ostrea edulis) (Cole 1942; Carriker 1955; Buhle and Ruesink 2009).

Food:

Oysters,Mussels, Barnacles

Trophic Status:

Carnivore

Carn

Habitats

General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatRockyNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone
Vertical HabitatLittoralNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)-1.5Field (Carriker 1955)
Maximum Temperature (ºC)30Hanks 1957, highest tested
Minimum Salinity (‰)12Minimum salinities survived by groups of Urosalpinx cinerea varied from 12 to 16 ppt for animals from Norfolk VA and Beaufort NC exposed to gradual salinity reduction (Federghi 1931).
Minimum Reproductive Temperature15Experimental (Ganaros 1958)
Maximum Reproductive Temperature30Highest tested (Ganaros 1958)
Minimum Reproductive Salinity20Manzi 1970, experimental
Minimum Length (mm)16.5Sauber 1943, cied by Carriker 1955
Maximum Length (mm)35Excluding localized giant populations (eg. , Virginia Atlantic bays), which can reach 55-61 mm length
Broad Temperature RangeNoneCold temperate-Subtropical
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

Urosalpinx cinerea is a major predator on barnacles and bivalves in low-intertidal and shallow subtidal waters. Its predation on oysters is especially ecologically and economically important, and it is widely regarded as a serious pest in oyster fisheries and in restoration of oyster populations (Cole 1942; Carriker 1955; Buhle and Ruesink 2009; Kimbro et al. 2009).

Economic impacts

Fisheries: Urosalpinx cinerea is regarded as a serious pest to oyster culture and fisheries in its native East Coast waters, and in many of the bays to which it has been introduced. The activities that oyster growers perform to encourage oysters, including planting 'cultch' (shells to promote settlement) and transplanting oysters to new grounds, tend to spread the Oyster Drill (Carriker 1955). The effects of its predation are often-underestimated, since small spat are the most likely to be eaten, affecting recruitment (Cole 1942). A variety of methods are used to control them, including hand-picking of snails and egg-cases, using special dredges with fine meshes, using screens to remove drills from dredged oysters, suction dredges to remove the drills after oysters are harvested, and traps that drills will crawl into for egg-laying (Cole 1942; Carriker 1955). Ironically, impacts of this drill in English waters, and probably in US waters, have been greatly reduced in recent decades, by reproductive abnormalities (imposex) caused by tributytin (TBT) used in antifouling paints. However, this pollutant also produces abnormalities in oysters and other mollusks so it is not a desirable control method (Gibbs et al. 1991; Mann et al. 2006).

Ecological Impacts

Predation: Urosalpinx cinerea is a predator on barnacles, mussels, oysters, and other bivalves, but only predatory impacts on oysters have been well-studied (Cole 1942; Carriker 1955; Buhle and Ruesink 2009; Kimbro et al. 2009). Aside from their commercial value, oyster beds are important as habitat and a food source for fishes and invertebrates, and for maintenance of water quality through filtration of the water column (e.g., Kennedy 1995; Newell et al. 2007). Predation on the Olympic Oyster (Ostrea lurida), the native oyster of the West Coast, in Tomales and Willapa Bays, is of particular concern, because it complicates attempts to restore this oyster, which is severely affected by over-harvesting, Pacific Oyster aquaculture, pollution, and shoreline development (Buhle and Ruesink 2009; Kimbro et al. 2009).

Competition: Urosalpinx cinerea is a potential competitor with native predatory snails. In estuaries in Essex and Kent, England, it largely replaced the native drill (Ocenebra erinacea). However, this was believed to be largely due to the invader's better tolerance of low temperatures during several unusually severe winters (Cole 1942). In Tomales Bay, predation by native crabs (Cancer antennarius) in high-salinity waters appeared to be the major factor separating U. cinera from a native whelk (Acanthinucella spirata), confining the invader to the inner estuary (Kimbro et al. 2009). In Willapa Bay, U. cinerea was largely spatially segregated from the Japanese Oyster Drill (O. japonica), and was also concentrated in the inner estuary, because of a requirement for higher temperatures for reproduction (Buhle and Rusesink 2009).

Regional Impacts

P110Tomales BayEconomic ImpactFisheries
An area of bay bottom used to hold Eastern Oysters (Crassostrea virginica), for sale, was 'heavily infested' with Atlantic Oyster Drills (Carriker 1955). Predation on cultured Crassostrea gigas is likely, but not reported (Carriker 1955).
P110Tomales BayEcological ImpactPredation
Urosalpinx cinerea is a major predator of the native Olympia Oyster (Ostrea lurida) in Tomales Bay, together with the native Acanthinucella spirata (Angular Unicorn Whelk). In Tomales Bay, predation by the large native Pacific Rock Crab (Cancer productus) excludes U. cinerea from the outer Bay, which is dominated by the better-defended native whelk. The Atlantic Oyster Drill is most common in the inner Bay, where it tolerates the low salinity, better than the native crab and whelk. The introduced Green Crab (Carcinus maenas) is the major predator in the inner Bay, but it is less effective at eating the introduced drill than the native crab, so U. cinerea reaches high densities and causes high mortality of Olympia Oysters (Kimbro et al. 2009; Cheng and Grosholz 2016). Ocean acidification may increase the predatory impact of U. cinerea on O. lurida in Tomales Bay, because the oysters raised under acidified conditions were smaller and more vulnerable to the drills (Sanford et al. 2013). In another study, Urosalpinx cinerea was a major predator on Olympia Oysters (Ostrea lurida), especially on the eastern side of the Bay, when low salinities and high temperatures, reduced predation by native Rock Crabs (Cancer productus). Climate warming my increase predatpory impacts by this snail (Cheng et al. 2016). Native predatory snails (Acanthinucella spirata) appeared to have little impact on Olymbic Oyster populations in this tuday (Cheng and Grosholz 2016).
P090San Francisco BayEcological ImpactPredation
In San Francisco Bay, U. cinerea feeds largely on the barnacle Balanus glandula and the exotic clam Corbula amurensis (Cohen 2005).
NEP-VNorthern California to Mid Channel IslandsEconomic ImpactFisheries
An area of bay bottom used to hold Eastern Oysters (Crassostrea virginica), for sale, was 'heavily infested' with Atlantic Oyster Drills (Carriker 1955). Predation on cultured Crassostrea gigas is likely, but not reported (Carriker 1955).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactPredation
Urosalpinx cinerea is a major predator of the native Olympia Oyster (Ostrea lurida) in Tomales Bay, together with the native Acanthinucella spirata (Angular Unicorn Whelk). In Tomales Bay, predation by the large native Pacific Rock Crab (Cancer productus) excludes U. cinerea from the outer Bay, which is dominated by the better-defended native whelk. The Atlantic Oyster Drill is most common in the inner Bay, where it tolerates the low salinity, better than the native crab and whelk. The introduced Green Crab (Carcinus maenas) is the major predator in the inner Bay, but it is less effective at eating the introduced drill than the native crab, so U. cinerea reaches high densities and causes high mortality of Olympia Oysters (Kimbro et al. 2009). Ocean acidification may increase the predatory impact of U. cinerea on O. lurida in Tomales Bay, because the oysters raised under acidified conditions were smaller and more vulnerable to the drills (Sanford et al. 2013). In another study, Urosalpinx cinerea was a major predator on Olympia Oysters (Ostrea lurida), especially on the eastern side of the Bay, when low salinities and high temperatures, reduced predation by native Rock Crabs (Cancer productus). Climate warming my increase predatpory impacts by this snail (Cheng et al. 2016). Native predatory snails (Acanthinucella spirata) appeared to have little impact on Olymbic Oyster populations in this tuday (Cheng and Grosholz 2016).
NEP-IVPuget Sound to Northern CaliforniaEcological ImpactPredation
Urosalpinx cinerea in Willapa Bay, feeds on the introduced Pacific Oyster (Crassostrea gigas) and the native Olympic Oyster (Ostrea lurida), but prefers the Pacific Oyster. The Atlantic Oyster Drill is largely restricted to the warmer inner portion of the estuary, where aquaculture farms are absent, and feral Pacific Oysters grow in hummocks. Urosalpinx cinerea prefers smaller oysters of both species, so Olympia Oysters are more vulnerable, because of their smaller adult size. Olympic Oysters do not survive in areas where Pacific Oysters are cultured, because of competition, predation, disturbance, and removal from cultured Pacific Oysters, so they frequently occur in the parts of the Bay inhabited by U. cinerea. Buhle and Rusesink (2009) conclude that Olympic Oyster restoration may only be feasible where Pacific Oyster aquaculture and oyster drills are absent.
NEP-IVPuget Sound to Northern CaliforniaEconomic ImpactFisheries
In Willapa Bay, Urosalpinx cinerea is rare in the deeper, colder parts of the bay, where Pacific Oysters (Crassostrea gigas) are cultured. However, it is abundant in areas where the native Olympia Oyster (Ostrea lurida) is present, making restoration of native oyster populations difficult in this bay (Buhle and Ruesink 2009).
CACaliforniaEcological ImpactPredation
Urosalpinx cinerea is a major predator of the native Olympia Oyster (Ostrea lurida) in Tomales Bay, together with the native Acanthinucella spirata (Angular Unicorn Whelk). In Tomales Bay, predation by the large native Pacific Rock Crab (Cancer productus) excludes U. cinerea from the outer Bay, which is dominated by the better-defended native whelk. The Atlantic Oyster Drill is most common in the inner Bay, where it tolerates the low salinity, better than the native crab and whelk. The introduced Green Crab (Carcinus maenas) is the major predator in the inner Bay, but it is less effective at eating the introduced drill than the native crab, so U. cinerea reaches high densities and causes high mortality of Olympia Oysters (Kimbro et al. 2009). Ocean acidification may increase the predatory impact of U. cinerea on O. lurida in Tomales Bay, because the oysters raised under acidified conditions were smaller and more vulnerable to the drills (Sanford et al. 2013). In another study, Urosalpinx cinerea was a major predator on Olympia Oysters (Ostrea lurida), especially on the eastern side of the Bay, when low salinities and high temperatures, reduced predation by native Rock Crabs (Cancer productus). Climate warming my increase predatpory impacts by this snail (Cheng et al. 2016). Native predatory snails (Acanthinucella spirata) appeared to have little impact on Olymbic Oyster populations in this tuday (Cheng and Grosholz 2016)., In San Francisco Bay, U. cinerea feeds largely on the barnacle Balanus glandula and the exotic clam Corbula amurensis (Cohen 2005)., Urosalpinx cinerea is a major predator of the native Olympia Oyster (Ostrea lurida) in Tomales Bay, together with the native Acanthinucella spirata (Angular Unicorn Whelk). In Tomales Bay, predation by the large native Pacific Rock Crab (Cancer productus) excludes U. cinerea from the outer Bay, which is dominated by the better-defended native whelk. The Atlantic Oyster Drill is most common in the inner Bay, where it tolerates the low salinity, better than the native crab and whelk. The introduced Green Crab (Carcinus maenas) is the major predator in the inner Bay, but it is less effective at eating the introduced drill than the native crab, so U. cinerea reaches high densities and causes high mortality of Olympia Oysters (Kimbro et al. 2009; Cheng and Grosholz 2016). Ocean acidification may increase the predatory impact of U. cinerea on O. lurida in Tomales Bay, because the oysters raised under acidified conditions were smaller and more vulnerable to the drills (Sanford et al. 2013). In another study, Urosalpinx cinerea was a major predator on Olympia Oysters (Ostrea lurida), especially on the eastern side of the Bay, when low salinities and high temperatures, reduced predation by native Rock Crabs (Cancer productus). Climate warming my increase predatpory impacts by this snail (Cheng et al. 2016). Native predatory snails (Acanthinucella spirata) appeared to have little impact on Olymbic Oyster populations in this tuday (Cheng and Grosholz 2016).
CACaliforniaEconomic ImpactFisheries
An area of bay bottom used to hold Eastern Oysters (Crassostrea virginica), for sale, was 'heavily infested' with Atlantic Oyster Drills (Carriker 1955). Predation on cultured Crassostrea gigas is likely, but not reported (Carriker 1955)., An area of bay bottom used to hold Eastern Oysters (Crassostrea virginica), for sale, was 'heavily infested' with Atlantic Oyster Drills (Carriker 1955). Predation on cultured Crassostrea gigas is likely, but not reported (Carriker 1955).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
P040 Newport Bay 1957 Non-native Established
NEP-VI Pt. Conception to Southern Baja California 1957 Non-native Established
P080 Monterey Bay 1945 Non-native Unknown
NEP-IV Puget Sound to Northern California 1941 Non-native Established
P130 Humboldt Bay 1941 Non-native Established
P110 Tomales Bay 1935 Non-native Established
P090 San Francisco Bay 1890 Non-native Established
NEP-V Northern California to Mid Channel Islands 1890 Non-native Established

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
697045 H. McCully in litt., 1941, to G.D. Hanna, cited in Carlton 1979; R. Baxter in litt., 1969, to A.G. S 1940 Humboldt Bay General Location Non-native 40.7864 -124.1922
697620 Introduced Species Study 2005 2005-11-16 Alcatraz Non-native 37.8253 -122.4223
697996 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-24 Fruitvale Bridge, San Francisco Bay Non-native 37.7690 -122.2296
698403 Townsend 1893; Stearns 1894; Stearns 1900 1889 Belmont, San Francisco Bay Non-native 37.5521 -122.2249
698607 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-27 Coyote Point Marina, San Francisco Bay Non-native 37.5907 -122.3180
699395 Introduced Species Study 2005 2005-07-06 Coyote Point Non-native 37.5920 -122.3210
699401 Introduced Species Study 2010 2010-06-13 Coyote Point Non-native 37.5920 -122.3210
699455 Introduced Species Study 2005 2005-06-07 Fort Point Non-native 37.8095 -122.4761
699559 Introduced Species Study 2010 2010-05-31 Redwood Creek - Shipping Non-native 37.5120 -122.2109
699878 Introduced Species Study 2010 2010-05-31 Redwood Creek - Marina Non-native 37.5021 -122.2130
699879 Introduced Species Study 2005 2005-09-07 Redwood Creek - Marina Non-native 37.5021 -122.2130
699926 Introduced Species Study 2010 2010-05-31 Dumbarton Bridge Non-native 37.5070 -122.1168
699934 Introduced Species Study 2005 2005-09-07 Dumbarton Bridge Non-native 37.5070 -122.1168
700472 Introduced Species Study 2010 2010-06-01 Coyote Point Marina Non-native 37.5905 -122.3177
700482 Introduced Species Study 2005 2005-09-09 Coyote Point Marina Non-native 37.5905 -122.3177
700699 V.L. Human, in litt., 1972, in Carlton 1979 1940 Newport Bay Non-native 33.6092 -117.9067
700802 Introduced Species Study 2005 2005-09-09 Sea Plane Harbor Non-native 37.6349 -122.3848
700958 A.G. Smith, pers. comm., in Burch 1945_ 1945 Elkhorn Slough General Location Non-native 36.8086 -121.7856
701933 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-24 Coast Guard Island Marina, San Francisco Bay Non-native 37.7812 -122.2458
702145 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-24 San Leandro Marina, San Francisco Bay Non-native 37.6966 -122.1932
702301 Introduced Species Study 2005 2005-08-19 Ayala Cove Non-native 37.8680 -122.4350
702896 Boyd et al. 2002 (Humboldt Bay Report) 2002 Bracut Non-native 40.8313 -124.0845
703224 Introduced Species Study 2005 2005-06-09 Point Cavallo Non-native 37.8319 -122.4737
703901 Introduced Species Study 2010 2010-06-30 Rodeo Marina Non-native 38.0394 -122.2717
704113 Introduced Species Study 2005 2005-10-05 Coast Guard Island Non-native 37.7812 -122.2457
704278 Cohen et al. 2005 (SF Bay Area RAS) 2004 2004-05-23 Brisbane Lagoon, San Francisco Bay Non-native 37.6862 -122.3906
704406 Bonnot 1935a 1935 Tomales Bay Non-native 38.2100 -122.9400
760384 Carlton 1979 1892 San Francisco Bay Non-native 37.8494 -122.3681
760385 Stearns 1899; 1900 1898 "Alameda flats" (off Oakland) Non-native 37.7521 -122.2643
760386 Stearns 1900 1899 near Redwood City Non-native 37.5232 -122.1938
760387 Keep 1901 1901 "Alameda shore" Non-native 37.7585 -122.2632
760388 J.L. Nicholson, pers. comm. in Hanna 1939 1928 San Francisco Bay Non-native 37.8494 -122.3681
760389 Smith and Gordon 1948 1948 Elkhorn Slough General Location Non-native 36.8086 -121.7856
760390 Haydock 1964 1962 Double Point, northern section Non-native 38.1228 -122.8635
760391 Haydock 1964 1962 Tomales Bay Oyster Company beds Non-native 38.1153 -122.8567
760392 Haydock 1964 1962 Abandoned oyster bed south of Millerton Point Non-native 38.1006 -122.8366
760393 Human 1971 1969 Upper Newport Bay Non-native 33.6344 -117.8895
760394 Wicksten 1978 1978 Coyote Point Non-native 37.5922 -122.3210
760395 Cohen and Chapman 2005 2005 2005-11-27 Dumbarton Bridge (pylon) Non-native 37.5031 -122.1230
768430 Ruiz et al., 2015 2013 2013-08-21 San Leandro Marina, San Francisco Bay, CA, California, USA Non-native 37.6980 -122.1908
771751 Ruiz et al., 2021a 2018 2018-09-19 Lake Merritt Boat House, San Francisco Bay, California, USA Non-native 37.8047 -122.2573
775501 Ruiz et al., 2022 2015 2015-09-24 Oakland Yacht Club, San Francisco Bay, California, USA Non-native 37.7839 -122.2641

References

Shalovenko, N. N. (2020) Tendencies of Invasion of Alien Zoobenthic Species into the Black Sea, Russian Journal of Biological Invasions 11(2): . 164–171

Abbott, R. Tucker (1974) American Seashells, Van Nostrand Reinhold, New York. Pp. <missing location>

Academy of Natural Sciences of Philadelphia 2002-2024a Malacology Collection Search. <missing URL>



Baldwin, Andy; Leason, Diane (2016) Potential Ecological impacts of Emerald Ash Borer on Maryland's Eastern Shore, In: None(Eds.) None. , <missing place>. Pp. <missing location>

Boersma, P. D; Reichard, S. H.; Van Burne, A. N. (2006) <missing title>, Invasive species in the Pacific Northwest, Seattle WA. Pp. 1-285

Bousfield, E. L. (1960) Canadian Atlantic Sea Shells, In: (Eds.) . , Ottawa. Pp. <missing location>

Bousfield, E. L., Thomas, M.L. H. (1975) Postglacial changes in distribution of littoral marine invertebrates in the Canadian Atlantic region, Proceedings of the Nova Scotian Institute of Science 27: 47-60

Boyd, Milton J.; Mulligan, Tim J; Shaughnessy, Frank J. (2002) <missing title>, California Department of Fish and Game, Sacramento. Pp. 1-118

Buhle, Eric R.; Ruesink, Jennifer L. (2009) Impacts of invasive oyster drills on Olympia oyster (Ostrea lurida Carpenter 1864) recovery in Willapa Bay, Washington, United States, Journal of Shellfish Research 28(1): 87-96

Carl, G. Clifford; Guiguet, C. J. (1972) Alien animals in British Columbia., British Columbia Provincial Museum: Department of Recreation and Conservation: Handbook 14: 1-102

Carlton, James T. (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific Coast of North America., Ph.D. dissertation, University of California, Davis. Pp. 1-904

Carriker, Melbourne Romaine (1955) Critical review of biology and control of oyster drills Urosalpinx and Eupleura., United States Fish and Wildlife Service Special Scientific Report 149: 1-150

Cheng, Brian S.; Grosholz, Edwin D. (2016) Environmental stress mediates trophic cascade strength and resistance to invasion, Ecosphere 7(4): e01247

Cheng, Brian S.;; Blumenthal, Jeffrey; Chang, Andrew L.; Barley, Jordanna; Matthew C. Ferner, K; Nielsen, arina J.;Ruiz, Gregory M.; Zabin, Chela J. (2021) Severe introduced predator impacts despite attempted functional eradication, Biological Invasions Published online: <missing location>

Cohen, Andrew N. 2005-2024 Exotics Guide- Non-native species of the North American Pacific Coat. https://www.exoticsguide.org/



Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Cohen, Andrew N.; Chapman, John T. (2005) <missing title>, San Francisco Estuary Institute, San Francisco. Pp. <missing location>

Cole, H. A. (1942) The American Whelk-Tingle, Urosalpinx cinerea (Say) on British oyster-beds., Journal of the Marine Biological Association of the United Kingdom 25: 477-508

De Kay, James E. (1843) Mollusca, Vol. 5, Zoology of New-York., In: (Eds.) Zoology of New-York. , Albany. Pp. <missing location>

Eissinger, Ann (2009) <missing title>, Nahkeeta Northwest Wildlife Services, Bow WA. Pp. 1-26

Eno, N. Clare (1996) Non-native marine species in British waters: effects and controls, Aquatic Conservation: Marine and Freshwater Ecosystems 6: 215-228

Eno, N. Clare; Clark, Robin A.; Sanderson, William G. (1997) <missing title>, Joint Nature Conservation Committee, Peterborough. Pp. <missing location>

Eno, N.C., Clark, R.A., Sanderson, W.G. 1997-2012 Directory of Non-Native Marine Species in British waters. <missing URL>



Faasse, Marco; Ligthart, Marianne (2009) American (Urosalpinx cinerea) and Japanese oyster drill (Ocinebrellus inornatus) (Gastropoda: Muricidae) flourish near shellfish culture plots in The Netherlands, Aquatic Invasions 4(2): 321-326

Faasse, Marco; Ligthart, Marianne (2007) The American oyster drill, Urosalpinx cinerea (Say, 1822), introduced to The Netherlands: increased risks after ban on TBT?, Aquatic Invasions 2(4): 402-406

Federighi, H. (1931) Salinity death-points of the oyster drill snail, Urosalpinx cinerea Say, Ecology 12(2): 346-353

Foss, Stephen (2009) <missing title>, California Department of Fish and Game, Sacramento CA. Pp. <missing location>

Foss, Stephen (2011) <missing title>, California Department of Fish and Game, Office of Spill Prevention and Response, Sacramento. Pp. 54

Galtsoff, Paul S. (1932) Introduction of Japanese Oysters into the United States, United States Department of Commerce, Bureau of Fisheries, Fisheries Circular 12: 1-16

Galtsoff, Paul S. (1964) The American oyster, Crassostrea virginica Gmelin. Chapter 18: Environmental factors affecting oyster populations, Fisheries Bulletin 64: 397-456

Ganaros, Anthony E. (1958) On development of early stages of Urosalpinx cinerea (Say) at constant temperatures and their tolerance to low temperatures., Biological Bulletin 114(2): 188-195

Gibbs, P. E.; Spencer, B. E.; Pascoe, P. l. (1991) The American oyster drill, Urosalpinx cinerea (Gastropoda): Evidence of decline in an imposex affected population (R. Blackwater, Essex), Journal of the Marine Biological Association of the United Kingdom 71: 827-838

Gosner, Kenneth L. (1978) A field guide to the Atlantic seashore., In: (Eds.) . , Boston. Pp. <missing location>

Gould, Augustus A. (1841) <missing title>, Folsom, Wells, and Thurston, Cambridge. Pp. <missing location>

Goulletquer, Philippe; Bachelet, Guy; Sauriau, Pierre; Noel, Pierre (2002) Invasive aquatic species of Europe: Distribution, impacts, and management, Kluwer Academic Publishers, Dordrecht. Pp. 276-290

Grason, Emily W.; Miner, Benjamin G. (2012b) Behavioral plasticity in an invaded system: non-native whelks recognize risk from native crabs, Oecologia 169: 105-115

Hanks, James E. (1957) The rate of feeding of the common oyster drill, Urosalpinx cinerea (Say), at controlled water temperatures, Biological Bulletin 112(3): 330-335

Hanna, G. Dallas (1966) Introduced mollusks of Western North America, Occasional Papers of the California Academy of Sciences 48: <missing location>

Harding, Julie M.; Mann, Roger (1999) Habitat and prey preferences of Veined Rapa Whelks (Rapana venosa) in the Chesapeake Bay: direct and indirect trophic consequences, Journal of Shellfish Research <missing volume>: 291

Hopkins, Dale R. (1986) Atlas of the distributions and abundances of common benthic species in San Francisco Bay, California, US Geological Survey Water Resources Investigations Report 86-4003: 1-16+ 25+228

Kennedy, Victor S. (1995) Ecological role of the eastern oyster, Crassostrea virginica, with remarks on disease., Journal of Shellfish Research 15(1): 177-183

Kim, Daemin; Taylor, Andrew T.; Near, Thomas J. (2022) Phylogenomics and species delimitation of the economically important Black Basses (Micropterus), Scientific Reports 12(9113): Published online
https://doi.org/10.1038/s41598-022-11743-2

Kimbro, David L. and 6 authors (2009) Invasive species cause large-scale loss of native California oyster habitat by disrupting trophic cascades., Oecologia 160: 563-575

Leidenberger, Sonja; Obst, Matthias; Kulawik, Robert; Stelzer, Kerstin; Heyer, Karin; Hardisty, Alex; Bourlat, Sarah J. (2015) Evaluating the potential of ecological niche modelling as a component in marine non-indigenous species risk assessments, Marine Pollution Bulletin 97: 470-487

Lippson, Alice Jane; Lippson, Robert L. (1997) <missing title>, Johns Hopkins University Press, Baltimore. Pp. <missing location>

Mann, Roger, Harding, Juliana M.; Westcott, Erica (2006) Occurrence of imposex and seasonal patterns of gametogenesis in the invading veined rapa whelk Rapana venosa from Chesapeake Bay., Marine Ecology Progress Series 310: 129-138

Manzi, John J. (1970) Combined effects of salinity and temperature on the feeding, reproductive, and survival rates of Eupleura caudata (Say) and Urosalpinx cinerea (Say) (Prosobranchia: Muricidae), Biological Bulletin 138(1): 35-46

McLean, James A. (2007) The Light and Smith Manual: Intertidal Invertebrates from Central California to Oregon, University of California Press, Berkeley CA. Pp. 713-1766

Mikkelsen, Paula M., Mikkelsen, Paul S., Karlen, David J. (1995) Molluscan biodiversity in the Indian River Lagoon, Florida, Bulletin of Marine Science 57(1): 94-127

Newell, Roger I. E. (1988) Ecological changes in the Chesapeake Bay: are they the result of overharvesting the American oyster, Crassostrea virginica?., In: (Eds.) Understanding the Estuary: Advances in Chesapeake Bay Research. Proceedings of a Conference.. , Baltimore, MD. Pp. <missing location>

Newell, Roger I. E.; Kennedy, Victor S.; Shaw, Kristi S. (2007) Comparative vulnerability to predators, and induced defense responses, of eastern oysters Crassostrea virginica and non-native Crassostrea ariakensis oysters in Chesapeake Bay., Marine Biology 152: 449-460

Pielou, E. C. (1991) <missing title>, University of Chicago Press, Chicago IL. Pp. 366

Quayle, D. B. (1969) Pacific oyster culture in British Columbia, Canadian Fisheries Research Board Bulletin 169: 1-192

Riggs, Sharon R. (2011) <missing title>, Padilla Bay NERR, Padilla Bay WA. Pp. 5

Robinson, John D.; Dillon, Robert T., Jr. (2008) Genetic divergence among sympatric populations of three species of oyster drills (Urosalpinx) in Cedar Key, Florida, Bulletin of Marine Science 82(1): 19-31

Rosenberg, Gary 1995-2023 Malacolog 4.1. http://www.malacolog.org/



Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>

Sanford, Eric and 5 authors (2013) Ocean acidification increases the vulnerability of native oysters to predation by invasive snails, Proceedings of the Royal Society of London B Published online: <missing location>

Say, Thomas (1822) An account of some of the marine shells of the United States, Journal of the Academy of Natural Sciences of Philadelphia 2: 221-248, 258-276,

Van den Brink, A. M.; Wijsman, J. W. M. (2010) <missing title>, ]IMARES - Institute for Marine Resources & Ecosystem Studies], Netherlands. Pp. 1-47

Verrill, A.E.; Smith, S.I. (1873) <missing title>, 1 Report of the United States Commission of Fish and Fisheries, <missing place>. Pp. 1-757

Walker, Randal L. (1971) Distribution of oyster drills, Urosalpinx cinerea in Wassaw Sound, Georgia, Georgia Journal of Science 39: 127-139

Wasson, Kerstin; Zabin, C. J.; Bedinger, L.; Diaz, M. C.; Pearse J. S. (2001) Biological invasions of estuaries without international shipping: the importance of intraregional transport, Biological Conservation 102: 143-153