Urosalpinx cinerea

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

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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).

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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.

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Taxonomy

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

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Habitats

General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
Salinity Range	Mesohaline	5-18 PSU
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
Vertical Habitat	Littoral	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-1.5	Field (Carriker 1955)
Maximum Temperature (ºC)	30	Hanks 1957, highest tested
Minimum Salinity (‰)	12	Minimum 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 Temperature	15	Experimental (Ganaros 1958)
Maximum Reproductive Temperature	30	Highest tested (Ganaros 1958)
Minimum Reproductive Salinity	20	Manzi 1970, experimental
Minimum Length (mm)	16.5	Sauber 1943, cied by Carriker 1955
Maximum Length (mm)	35	Excluding localized giant populations (eg. , Virginia Atlantic bays), which can reach 55-61 mm length
Broad Temperature Range	None	Cold temperate-Subtropical
Broad Salinity Range	None	Mesohaline-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).

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Regional Impacts

P110	Tomales Bay		Economic Impact	Fisheries
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).
NEA-II	None		Ecological Impact	Competition
Since its discovery in estuaries of Essex and Kent, U. cinerea largely replaced the native oyster drill (Ocenebra erinacea) in the 1930s-50s. Some of the American drill's success was apparently due to its greater tolerance of severe winters, which killed many of the native snails (Cole 1942). Now, both species are rare because of imposex, caused by tributlytin (TBT) pollution (Gibbs et al. 1991).
NEA-II	None		Economic Impact	Fisheries
Urosalpinx cinerea was regarded as a major pest when it began infesting beds of the European Oyster (Ostrea edulis) in England. Its biggest effect was on recruitment, since U. cinerea mostly consumes spat.  Fishermen used a variety of methods, from hand-picking and dredging, in an attempt to limit drill predation (Cole 1942).
NEA-II	None		Ecological Impact	Predation
Urosalpinx cinerea, in English oyster-beds, fed mostly on spat of European Oysters (Ostrea edulis). Spat of about 10 cm were preferred to smaller spat, adult oysters, or to mussels (Cole 1942).
P110	Tomales Bay		Ecological Impact	Predation
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).
P090	San Francisco Bay		Ecological Impact	Predation
In San Francisco Bay, U. cinerea feeds largely on the barnacle Balanus glandula and the exotic clam Corbula amurensis (Cohen 2005).
P270	Willapa Bay		Ecological Impact	Predation
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 aquaculure 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 be feasible only where oyster aquaculture and oyster drills are absent.
P270	Willapa Bay		Economic Impact	Fisheries
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).
NEP-V	Northern California to Mid Channel Islands		Economic Impact	Fisheries
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-V	Northern California to Mid Channel Islands		Ecological Impact	Predation
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-IV	Puget Sound to Northern California		Ecological Impact	Predation
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-IV	Puget Sound to Northern California		Economic Impact	Fisheries
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).
CA	California		Ecological Impact	Predation
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).
CA	California		Economic Impact	Fisheries
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).
WA	Washington		Ecological Impact	Predation
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 aquaculure 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 be feasible only where oyster aquaculture and oyster drills are absent.
WA	Washington		Economic Impact	Fisheries
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).

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