Description
The initial identification of Rapana venosa from Chesapeake Bay was confirmed by: Dr. Yuri Kantor, Russian Academy of Sciences, Moscow, and Dr. M. Gerry Harasewych, Curator of Molluscs, Smithsonian Institution, Washington D.C.
Synonymy, Potentially Misidentified species- Black Sea populations of Rapana venosa have been frequently referred to as R. thomasiana, a synonym, and R. bezoar, another species, native to the South China Sea (Virginia Institute of Marine Science 1998; Mann and Harding 2000).
Taxonomy
Kingdom | Phylum | Class | Order | Family | Genus |
---|---|---|---|---|---|
Animalia | Mollusca | Gastropoda | Stenoglossa | Muricidae | Rapana |
Synonyms
Invasion History
Chesapeake Bay Status
First Record | Population | Range | Introduction | Residency | Source Region | Native Region | Vectors |
---|---|---|---|---|---|---|---|
1998 | Established | Expanding | Introduced | Regular Resident | Eastern Atlantic | Western Pacific | Shipping(Ballast Water, Fouling Community) |
History of Spread
Rapana venosa (Veined Rapa Whelk) is native to the Northeast Pacific, from the southern Pacific coast of Russia, to the Sea of Japan, Yellow Sea, and East China Sea (Golikov et al. 1976; Mann and Harding 2000). In 1947, it was first collected from the Black Sea, in Novorossiyski Bay, where it was probably transported by ships, or with Crassostrea gigas (Pacific Oysters). Within a decade, it dispersed widely in the Black Sea and entered the sea of Azov (Chukhchin 1984; Zolotarev 1996; Mann and Harding 2000).
Between 1977 and 1984, it colonized the westernmost Black Sea, and the Sea of Marmara (Turkey) (Kinzelbach 1986).
The spread of R. venosa into the Mediterranean is patchy, suggestive of jumps involving long-range ship dispersal, followed by larval dispersal over shorter distances. In 1974, Rapana venosa was first collected in the Adriatic Sea, where it was probably transported by ship, reaching Venice by 1983 (Cezari and Pelizzato 1985). In in 1980, it was collected off Alba, on the West side of Italy, and in 1986, the Aegean Sea to the East (Koutsoubas and Voultsiadou-Koukoura 1990).
A single specimen of Rapana venosa has been collected from England in 1992 (Eno et al. 1997), as well as several specimens from Brittany, France (Mann and Harding 2000). Two specimens were collected in North Sea in the UK and Netherlands offshore waters 2005 (Kerckhoff et al. 2006). Rapana venosa has had scattered occurrences on the Atlantic coast of Europe, and has at least one established population. A single specimen of Rapana venosa has been collected from England in 1992 (Eno et al. 1997; (Mann and Harding 2000). One specimen was found in the Gulf of Morbihan, Brittany in 1998, and a small but stable population is established in the Bay of Quiberon (Goulletquer et al. 2002; Kerckhof et al. 2006; Chandler et al. 2008). A single specimen was collected in the Ría de Arousa, Galicia, Spain in 2007 (Rolan and Banon-Diaz 2007; Banon et al. 2008). Two specimens were collected in the North Sea in UK and Netherlands offshore waters in 2005 (Kerckhoff et al. 2006). A few specimens were found on transplanted Crassostrea gigas in Puget Sound and Willapa Bay, but these populations did not become established (Hanna 1966; Carlton 1979) populations did not become established. In 1999, an adult R. venosa and egg capsules were collected at the mouth of the La Plata estuary in Argentina. This whelk is established and abundanr on this part of the South American coast (Pastorino et al. 2000). Giberto et al. 2006; Lanfranconi et al. 2009; Carranza et al. 2010).
In June 1998, 2 specimens of R. venosa were collected in the lower James River. Portsmouth VA. Collection notes are given below:
'Specimens collected in the lower James River:
Date: June 5 1998
Method of collection: 30' otter trawl
Collected by: Patrick Geer and Michael Land, VIMS Trawl Survey Group. Location: Latitude 36o 57.12' N, Longitude 76o 24.86' W.
VIMS Trawl survey cruise number JA 980605 station #6. This is below the I-664 bridge tunnel in Hampton Roads, south west of the coal terminal piers, on the south side of the river channel. Two specimens, one is in alcohol, went to the Smithsonian and back, and is in the invert collection at VIMS.' (Roger Mann 1998).
Subsequently, Rapana venosa was found to have a well-established population in lower western Chesapeake Bay, occurring from the Bay Bridge-Tunnel to the mouth of the Rappahannock River and the opposite Eastern Shore. More than 1000 individuals have been collected, as well as numerous egg-cases (Harding and Mannn 1999; Mann and Harding 2000). The expected range of R. venosa within the Bay extends at least to the northern mouth of the Rappahannock River (1 specimen, 1998) (Mann and Harding 2000), and perhaps further up the Bay, since adults tolerate salinities of 9-12 ppt in the Sea of Azov (Chukhchin 1984). Since 2003, three range extensions of R. venosa have been reported with the Bay: (1) seaward along the southern edge of the Bay to Cape Henry (2): upstream in the James River about 10 km, from the VA 258 bridge to Thomas Rock and Brown Shoal (oysterbeds); (3) northward
; northward in the Bay to Tangier Light (Harding and Mann 2005)
Based on temperature tolerance, Rapana venosa's future Atlantic coast range could extend from Cape Cod to Cape Hatteras NC or Charleston SC. The long planktonic period of larvae (14-17 days), and the potential for coastwise ship transport suggest that rapid coastal range expansion is possible (Mann and Harding 2000). Juvenile R. venosa have been found attached to Loggerhead Sea Turtles (Caretta caretta) in Wassaw Sound GA, so populations may exist outside the Chesapeake Bay (Harding et al. 2011).
History References- Carlton 1979; Cezari and Pelizzato 1985; Chukhchin 1984; Eno et al. 1997; Golikov 1976; Hanna 1966; Harding and Mann 1999; Harding and Mann 2005; Kerckhoff et al. 2006; Kinzelbach 1986; Koutsoubas and Voultsiadou-Koukoura 1990; Mann and Harding 2000; Roger Mann 1998; Pastorino et al. 2000; Virginia Institute of Marine Science 1998; Zolotarev 1996
Invasion Comments
Ecology
Environmental Tolerances
For Survival | For Reproduction | |||
---|---|---|---|---|
Minimum | Maximum | Minimum | Maximum | |
Temperature (ºC) | 4.0 | 35.0 | 18.0 | 26.0 |
Salinity (‰) | 9.0 | 39.0 | 12.0 | |
Oxygen | hypoxic | |||
pH | ||||
Salinity Range | meso-eu |
Age and Growth
Male | Female | |
---|---|---|
Minimum Adult Size (mm) | 35.0 | 35.0 |
Typical Adult Size (mm) | 58.0 | 58.0 |
Maximum Adult Size (mm) | 78.0 | 78.0 |
Maximum Longevity (yrs) | ||
Typical Longevity (yrs |
Reproduction
Start | Peak | End | |
---|---|---|---|
Reproductive Season | |||
Typical Number of Young Per Reproductive Event |
|||
Sexuality Mode(s) | |||
Mode(s) of Asexual Reproduction |
|||
Fertilization Type(s) | |||
More than One Reproduction Event per Year |
|||
Reproductive Startegy | |||
Egg/Seed Form |
Impacts
Economic Impacts in Chesapeake Bay
Rapana venosa's (Veined Rapa Whelk's) invasion is expected to have economic impacts on shellfisheries in Chesapeake Bay because of its role as a a very efficient predator of bivalves. Populations of Mercenaria mercenaria (Hard Clams) are expected to be most affected. This fishery has annual landings of abot 1.1 million pounds, with a dockside value of $6 million (Harding and Mann 1999). The harvest is currently declining, due in part to overfishing, pollution, and disease (Savini et al. 2002). Conseqently, the addition of a new predator is viewed with alarm. In laboratory experiments, small R. venosa (60-100 mm) consumed ~3.6% of their body weight in M. mercenaria per day, while large specimens (>101 mm) ate ~0.8% of their body weight per day (Savini et al. 2002). Savini et al. estimate that a population of 1,000 R. venosa could reduce the Hard Clam annual Hard Clam harvest by 0.3-0.9% per year. Thus, it appears likely that the present small population of R. venosa is having at least a small impact on the Hard Clam harvest. Predation on Crassostrea virginica (Eastern Oyster) is expected to be less significant, because most remaining beds are in low-salinity waters, near the tolerance limits of R. venosa (Mann and Harding 2000). References- Harding and Mann 1999; Mann and Harding 2000
Economic Impacts Outside of Chesapeake Bay
The invasion of the Black Sea by Rapana venosa (Veined Rapa Whelk) severely affected shellfisheries in the Black Sea, severely reducing populations of oysters (Ostrea edulis), scallops (Pecten ponticus), mussels (Mytilus galloprovincialis), and clams (Venus gallina) (Chukhchin 1984; Zolotarev 1996). Attempts to eradicate R. venosa on the Bulgarian Coast were unsuccessful (Staykov 1997).
In the Black Sea (e.g. Bulgaria, Turkey), commercial fisheries have developed for R. venosa, in which the snails are shipped to Japan and Korea. In Turkey, size limits have been imposed to conserve stocks (Alpbaz and Temelli 1997; Staykov 1997).
In the northwest Atlantic, Rapana venosa is expected to eventually extend its range, at least to Cape Hatteras in the South and Cape Cod in the north (Mann and Harding 2000).
References- Alpbaz and Temelli 1997; Chukhchin 1984; Mann and Harding 2000; Staykov 1997; Zolotarev 1996
Ecological Impacts on Chesapeake Native Species
At present, ecological impacts of Rapana venosa (Veined Rapa Whelk) in the Chesapeake Bay are still small, but experiences in the Black Sea and Mediterranean suggest that this snail could be a significant predator in the future (Chukhchin 1984; Zolotarev 1996; Harding and Mann 1999).
Predation- In the Black Sea, this mollusk: 'Feeds primarily on bivalve mollusks, paralyzing them with toxins and eating them with the aid of its soft proboscis. In aquariums, Rapana venosa eats Mytilus (mussels), Ostrea (oysters), Tapes (clams), Venus(clams), Pecten (scallops), and Cardium (cockles), and the gastropod mollusk Patella (limpets). When Rapana is offered mussels and oysters simultaneously, it clearly prefers the former. This is explained, probably, by the thinner shell of the mussels which Rapana can more easily penetrate. Young-of-the year Rapana venosa eat Balanus improvisus. Rapana venosa may also feed on carrion. In the aquarium, they eat the meat of mussels, oysters, dead fish and crabs.' (Chukhchin 1984).
'Before its establishment in the Black Sea, predatory gastropods played an insignificant role in the shellfish bed biocoenosis. The Black Sea lacked predatory marine organisms, including predatory gastropod mollusks which could feed on bivalves. The establishment of Rapana created a new ecological niche. This brought about substantial changes in the biocoenosis of the Black Sea shellfish beds- one dominant species replaced others. In the Gudaytskii oyster bank, before the establishment of Rapana, 4 species of lamellibranch bivalves predominated, Ostrea taurica, Mytilus galloprovincialis, Pecten ponticus, and Modiola adriatica (Nikitin 1934). Our studies on the Gudaytskii oyster bank showed that in 1958, after the full destruction of large mollusks by Rapana, the dominant role went to the small mollusk Gouldia minima (Chukhchin 1960).'
It should be noted that in the Black Sea, prior to R. venosa's invasion, there were no large predatory gastropods, while several native species occur in the Chesapeake Bay and northwest Atlantic.
In the Chesapeake Bay, Rapana venosa in aquaria consumed Mercenaria mercenaria (Hard Clams), but are likely to feed on other bivalves as well. The extent of predation in the field is difficult to determine, because R.venosa does not leave a clear signature, such as boring or rasping marks on bivalve shells (Harding and Mann 1999; Mann and Harding 2000). Nearly all collections have been from soft bottoms, in the lower Bay, where the dominant large bivalve is M. mercenaria. Impacts on Crassostrea virginica (Eastern Oysters) are expected to be small, because diseases have confined substantial oyster populations to low salinities (Mann and Harding 2000).
In laboratory experiments, small R. venosa (60-100 mm) consumed ~3.6% of their body weight in M. mercenaria per day, while large specimens (>101 mm) ate ~0.8% of their body weight per day (Savini et al. 2002). Savini et al. estimate that a population of 1,000 R. venosa could reduce the Hard Clam annual Hard Clam harvest by 0.3-0.9% per year.
Competition- Other predatory gastropods with similar feeding habits are Busycotypicus caniculatus (Channeled Whelk), Busycon carica (Knobbed Whelk), B. sinistrum (Lightning Whelk), and Neverita duplicata (Shark Eye Snail). Competition between these species and Rapana venosa has not been studied.
Habitat Change- One effect that R. venosa's invasion may have involves changing the shell resources for hermit crabs, possibly favoring the now-rare Clibanarius vittatus (Striped Hermit Crab) over the dominant Pagurus pollicaris (Flat-Clawed Hermit Crab). Rapana venosa's morphology appears better suited to the former crab (Harding and Mann 1999).
References- Chukhchin 1960; Harding and Mann 1999; Mann and Harding 2000; Savini et al. 2002; Zolotarev 1996
Ecological Impacts on Other Chesapeake Non-Native Species
Impacts of Rapana venosa(Veined Rapa Whelk) on introduced organisms in the Chesapeake Bay are not known. However, if exotic oysters such as Crassostrea ariakensis (Suminoe Oyster) are introduced, R. venosa is a potential predator.
References
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