Rapana venosa (veined rapana whelk)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Social Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Rapana venosa (Valenciennes, 1846)
Preferred Common Name
- veined rapana whelk
Other Scientific Names
- Rapana fusiformis Martens, 1902
- Rapana pontica Nordsieck, 1968
- Rapana thomasiana Crosse, 1861
International Common Names
- English: Asian rapa whelk
Local Common Names
- Italy: bobolone; cocozza
- Japan: aka-nishi
- Netherlands: geaderde stekelhoorn
Summary of InvasivenessTop of page
R. venosa is considered as one of worst invaders worldwide. It has a high ecological fitness as evidenced by its high fertility, fast growth rate and broad tolerance to salinity, temperatures, water pollution and oxygen deficiency, giving it all the characteristics of a successful invader (Kerckhof et al., 2006). In areas where it has been introduced it has caused significant changes to the ecosystem (ISSG, 2007).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Mollusca
- Class: Gastropoda
- Subclass: Caenogastropoda
- Order: Neogastropoda
- Unknown: Muricoidea
- Family: Muricidae
- Genus: Rapana
- Species: Rapana venosa
Notes on Taxonomy and NomenclatureTop of page
Rapana venosa is a gastropod mollusc in the family Muricidae which is placed in the clade Neogastropoda of the Caenogastropoda according to the classification of Bouchet and Rocroi (2005).
DescriptionTop of page
Adult R. venosa have a heavy globose shell with a large inflated body whorl, a deep umbilicus, and a low conical spire. The columella is broad, smooth, and slightly concave, and the aperture is large, ovate, and slightly expanded with a thin outer lip and the edge fluted to match the external spirals. Small elongate teeth are present along the edge of the outer lip. The spiral sculpture is irregular with the external shell ornamentation including smooth spiral ribs that end in regular blunt knobs at both the shoulder and the periphery of the body whorl. In addition, fine spiral ridges are crossed by low vertical riblets. The siphonal canal is broad, widely open, and bearing a series of scales.
DistributionTop of page
The native worldwide distribution of R. venosa includes the temperate Sea of Japan, the Yellow Sea, the Bohai Sea, the East China Sea to Taiwan in the south, and Peter the Great Bay off Vladivostok in the north (Mann and Harding, 2003). Currently, there are five known geographic regions containing reproducing populations of R. venosa that are distinct from the native (Asian) population (ICES, 2004). These are the Black Sea, the Adriatic and Aegean Seas, Chesapeake Bay in the northwest Atlantic, the Rio de la Plata Estuary in the southwest Atlantic, and the coast of Brittany in France in the northeast Atlantic (ICES, 2004). Limited records have also been made on the Pacific coast of Canada and in Willapa Bay, Washington, USA, where this species is not considered to be established (Mann and Harding, 2003).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Atlantic, Northeast||Localised||Introduced||ICES, 2004||Bay of Quiberon, France|
|Atlantic, Northwest||Localised||Introduced||Invasive||ICES, 2004||Chesapeake Bay. VA|
|Atlantic, Southwest||Localised||Introduced||Giberto et al., 2004||Rio de la Plata estuary|
|Mediterranean and Black Sea||Widespread||Introduced||Invasive||ICES, 2004|
|Pacific, Eastern Central||Unconfirmed record||Introduced||ICES, 2004||Willapa Bay, WA|
|Pacific, Northwest||Widespread||Native||Not invasive||ICES, 2004|
|Pacific, Southwest||Unconfirmed record||Introduced||ICES, 2004|
|Pacific, Western Central||Widespread||Native||Not invasive||ICES, 2004|
|China||Present||Present based on regional distribution.|
|-Fujian||Widespread||Native||Not invasive||ICES, 2004|
|-Hebei||Widespread||Native||Not invasive||ICES, 2004|
|-Jiangsu||Widespread||Native||Not invasive||ICES, 2004|
|-Liaoning||Widespread||Native||Not invasive||ICES, 2004|
|-Shandong||Widespread||Native||Not invasive||ICES, 2004|
|-Tianjin||Widespread||Native||Not invasive||ICES, 2004|
|-Zhejiang||Widespread||Native||Not invasive||ICES, 2004|
|Japan||Present||Present based on regional distribution.|
|-Hokkaido||Widespread||Native||Not invasive||ICES, 2004|
|-Honshu||Widespread||Native||Not invasive||ICES, 2004|
|-Kyushu||Widespread||Native||Not invasive||ICES, 2004|
|-Ryukyu Archipelago||Widespread||Native||Not invasive||ICES, 2004|
|-Shikoku||Widespread||Native||Not invasive||ICES, 2004|
|Taiwan||Widespread||Native||Not invasive||ICES, 2004|
|USA||Present||Present based on regional distribution.|
|-Virginia||Localised||Introduced||1998||Invasive||ICES, 2004||Chesapeake Bay|
|-Washington||Unconfirmed record||Introduced||ICES, 2004||Willapa Bay|
|Argentina||Localised||Introduced||1998||Giberto et al., 2004||Rio de la plata Estuary|
|Uruguay||Localised||Introduced||1998||Giberto et al., 2004||Rio de la plata Estuary|
|Albania||Present||Introduced||2011||Ruci et al., 2014|
|Belgium||Present||2006||Kerckhof et al., 2006|
|France||Present||Introduced||1998||Goulletquer et al., 2002||Bay of Quiberon|
|Greece||Present||Introduced||1986||Koutsoubas and Voultsiadou-Koukoura, 1991||Thermaikos Gulf|
|Italy||Widespread||Introduced||1973||Not invasive||Ghisotti, 1974||Northern Adriatic; (Ionian: Crocetta and Soppelsa, 2006)|
|Netherlands||Present, few occurrences||Introduced||2005||Kerckhof et al., 2006||Off Scheveningen, accidental|
|Romania||Widespread||Introduced||1961||Invasive||Grossu, 1964||Black Sea coast|
|Russian Federation||Present||Present based on regional distribution.|
|-Russian Far East||Widespread||Native||ICES, 2004||Peter the Great Bay|
|-Southern Russia||Widespread||Introduced||1947||Invasive||Drapkin, 1963||Black Sea coast|
|Slovenia||Introduced||Min and Vio, 1997|
|Spain||Present||Introduced||2007||Rolán and Bañón, 2007|
|UK||Present||Introduced||2005||Kerckhof et al., 2006||30 km South of the Dogger Bank, record unconfirmed (1991); established (2005)|
|Ukraine||Widespread||Introduced||1954||Invasive||Chukhchin, 1961||Black Sea coast|
|New Zealand||Unconfirmed record||Introduced||ICES, 2004|
History of Introduction and SpreadTop of page
The introduction of R. venosa into the Black Sea is suspected to have occurred some time in the 1940s (probably 1947) (Saglam and Duzgunes, 2007), with the first record of observation being in Novorossiysky Bay in Russia (Drapkin, 1963). It was most probably introduced into the area as a species associated with oyster seed (Mann and Harding, 2003). The culture of Japanese oysters was recorded in this region in the early 1940s. Once established in a founder location, the invasion of the rest of the Black Sea could have been facilitated by planktonic larvae dispersal alone without the need to invoke other vectors. Range extension occurred along the Caucasian and Crimean coasts and to the Sea of Azov (1953) within a decade of the first report, and subsequently to the northwest Black Sea, and the coastlines of Romania (1961), Bulgaria (1955), Turkey (1959) and the Marmara Sea (1969) (Kaneva-Abadjieva, 1958; Fischer-Piette, 1960; Chukhchin 1961; Grossu, 1964).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Albania||Italy||2011||Ruci et al. (2014)|
|Argentina||1998||Giberto et al. (2004)|
|Belgium||2006||Kerckhof et al. (2006)|
|France||Italy||1998||Goulletquer et al. (2002)|
|Greece||1986||Koutsoubas and Voultsiadou-Koukoura (1991)||Introduced from the Black Sea|
|Russian Federation||Japan||1947||Drapkin (1963)|
|Slovenia||Italy||1997||Min and Vio (1997)|
|Turkey||1960||Fischer-Piette (1960)||Introduced from the Black Sea|
|UK||2005||Kerckhof et al. (2006)|
|Uruguay||1998||Giberto et al. (2004)|
|USA||1988-1998||Harding and Mann (1999)|
Risk of IntroductionTop of page
The life history of R. venosa makes it a viable candidate for continuing range expansion and new invasions facilitated by ballast water vectors (ICES, 2004). Furthermore, the modern global economy and associated human vectors for transport across oceans provide continuous means of introduction of this robust animal into new receptor regions (ICES, 2004).
The emergence of ballast water as a major vector in marine introductions over the past half-century, combined with the imposing volumes of water moved as part of commercial ship traffic, suggest that continued dispersal of R. venosa will occur within the native and established introduced ranges as well as in new regions (ICES, 2004).
Establishment over a period of decades by natural dispersal in estuaries and coastal regions from Cape Cod to Cape Hatteras is considered a high probability. Observations on R. venosa biology and physiological tolerances in the Chesapeake Bay strongly suggest that this animal is capable of successful colonization and establishment of viable populations within estuarine habitats up and down the east coast of the USA (Mann and Harding, 2003). Its dispersal rate may be enhanced in that Hampton Roads Virginia, an area a very close proximity of the Chesapeake Bay, serves as a major container port for shipping along the Atlantic coast and in trade with Europe and Asian ports (ICES, 2004).
In both Chesapeake Bay and Rio de la Plata, one or more major ports exist that could, through ballast water, serve as potential donor locations to support new introductions (ICES, 2004). As far as the westward movement of R. venosa from the Black Sea towards the Mediterranean is concerned, the current progression rate is slow but expected to continue in the future (ICES, 2004).
Plasticity in the duration of the pelagic larval phase, the absence of specific larval settlement cues, broad dietary options in the early post-settlement stage, rapid growth to possible predation refuge, relatively early onset of sexual maturity, high fecundity, considerable longevity, and tolerance of challenging environments with respect to anthropogenic stressors make R. venosa a formidable invasion threat to regions that are both served by the identified vectors and within the broad niche descriptors (ICES, 2004).
HabitatTop of page
R. venosa is a habitat generalist and exploits practically every available prey. It occupies an empty ecological niche in the Black Sea and has exerted significant predatory pressure on the indigenous malacofauna.
Habitat ListTop of page
|Coastal areas||Principal habitat||Natural|
|Estuaries||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Inshore marine||Principal habitat||Natural|
|Benthic zone||Secondary/tolerated habitat||Natural|
Biology and EcologyTop of page
In order to provide guidelines for fisheries management, the genetics population structure was assessed using 10 polymorphic allozyme loci from seven populations of R. venosa across the species’ range on the Chinese coast. Significant genetic differentiation was present, and the h value was 0.016 across all populations (Yang et al., 2008).
A total of 57 polymorphic microsatellite loci were developed for R. venosa through 454 sequencing. The microsatellite loci described in this study will facilitate investigation of conservation genetics of this species (Sun et al., 2014).
The isolation, identification and characterisation of 11 novel antimicrobial peptides produced by the haemolymph of R. venosa was explored. The haemolymph was isolated from the foot of the animal. Four of the Pro-rich peptides showed strong antimicrobial activities against tested microorganisms including Gram-positive and Gram-negative bacteria (Dolashka et al., 2011).
As in most prosobranchs the sexes are separate. The gonad is situated dorsal to and in close contact with the digestive gland. Fertilization is internal and eggs are deposited in well-defined capsules that are formed in the oviduct before transferring into the egg capsule gland where they become visible. After leaving the oviduct, the egg capsule, which quickly hardens on contact with seawater, is generally transferred to the foot for deposition. These egg masses are then fixed to the benthic substrata such as mollusc shells and /or rocks (Webber, 1977 in Saglam and Duzgunes, 2007).
Clusters of these egg cases about 30 mm high are individually attached to the substrate and larvae hatch from there after 12-17 days and remain another 14-17 days in the plankton (Zenetos et al., 2004). Metamorphosis of the larvae is easily defined by the loss of the larval velum, an organ responsible for feeding, swimming and gas exchange. This transformation marks the transition from a swimming planktonic stage to a largely sedentary benthic stage (Zimmerman and Pechenik, 1991 in Saglam and Duzgunes, 2007). All larval stages exhibit 48 hour tolerance to salinities as low as 15 ppt with minimal mortality. The pattern of larval development and the identification of the distinct stages of veliger larvae are described by Saglam and Duzgunes (2007). Recently settled individuals grow at >1 mm per week, reaching shell lengths of 40-50 mm within five months post-settlement and >60 mm at 1 year (ICES, 2004).
In its native range mating of R. venosa occurs over an extended period during the winter and spring preceding egg-laying (ICES, 2004). The laying of eggs takes place from April to late July, a period that corresponds to a temperature range of 13°C to 26°C (ICES, 2004). One female adult can lay multiple egg masses throughout the course of one summer without intervening mating events. Significant plasticity in the duration of the planktonic phase has been observed, with metamorphosis being observed as late as 80 days after hatching (ICES, 2004).
In the Black Sea the observed reproductive period is July to September (with a temperature window of 19 to 25°C), whereas in eastern regions a spawning period of May to November has been reported (ICES, 2004). In the northern Adriatic, the deposition of egg masses takes place from the end of March to mid-September (ICES, 2004).
Growth is rapid during the first year of life (20 to 40 mm) and maturation/reproduction occurs from the second year onwards (Zenetos et al., 2004). R. venosa adults display mating activity all year in laboratory populations (Mann and Harding, 2003).
In in vitro experiments by Saglam and Duzgunes (2007), the number of egg capsules per egg mass ranged from 3 to 363 with a mean of 84. The total number of egg capsules per adult female in the spawning season of different sizes of R. venosa ranged from 197 to 999 with a mean of 575. Capsule size was strongly correlated with female size, but the number of egg capsules laid per female was independent of female size. Five different stages of development of veliger larvae in the capsule were identified. R. venosa demonstrated large numbers of embryos per egg mass, relatively small size of the egg, the absence of nurse cells and small size of the larva at its terminal stage of intracapsular development prior to hatching. The fecundity from each individual ranged from 109,000 to 555,000 eggs, values that are similar to those published by Chung et al. (2002) for R. venosa from Korea (Saglam and Duzgunes, 2007).
Imposex and decline in reproductive output in marine gastropods have been linked to tributyltin (TBT) exposure. In a study of R. venosa imposex incidence and sex ratios from 1998 to 2009 carried out at Chesapeake Bay, USA, tissue TBT concentrations (ng g-1) were examined with respect to whelk sex, size, and water temperature at the time of collection, and also to egg case size, hatching success, and veliger diameter. Imposex incidence declined and population sex ratios moved closer to parity from 1998 to 2009. Exponential declines in TBT concentrations from female-specific first to last clutches within a reproductive season were observed, indicating that whelks depurate TBT through egg case deposition. Egg capsule hatching success and veliger size were similar for female and imposex whelks. The R. venosa imposex levels observed in Chesapeake Bay apparently do not affect the production, release, or viability of larvae (Harding et al., 2013).
The occurrence and frequency of multiple paternity was studied by applying five highly polymorphic microsatellites in R. venosa in 19 broods (1381 embryos) collected from Dandong, China.(Xue et al., 2014). Results indicate that a high level of multiple paternity occurs in the wild population of R. venosa. Similar patterns of multiple paternity in the 2-6 assayed capsules from each brood imply that fertilization events within the body of a female occur mostly (but not entirely) as random draws from a “well-but-not-perfectly blended sperm pool” of her several mates. Strongly skewed distributions of fertilization success among sires also suggest that sperm competition and/or cryptic female choice might be important for post-copulatory paternity biasing in this species.
Physiology and Phenology
Populations on hard rock substrates have predominantly dark-coloured shells, whereas populations on adjacent sand exhibit a higher frequency of white or pale brown shell (ICES, 2004). Also, individuals from the northern Adriatic collected on sand substratum were significantly smaller than individuals collected on breakwaters. These hard substrates present valuable prey-rich resources for R. venosa, as demonstrated by the rapid invasion of the resource when resident adults are intentionally removed. Individuals in such locations have been observed feeding in temperatures as low as 8°C (ICES, 2004).
In planktonotrophic veligers, the most important food sources are flagellates, diatoms, organic and inorganic particles (Webber, 1977). Young R. venosa are generalist predators and consume large numbers of barnacles, mussels, oyster spat, and oysters (ICES, 2004). Adults feed on bivalves and may be scavengers on carrion. Harding and Mann (1999) reported that R. venosa can open a clam by smothering the shell and introducing the proboscis between the gaping valve, without any drilling (Zenetos et al., 2004).
Studies under field and laboratory conditions found that R. venosa are nocturnal and remain burrowed most of the day, avoiding settlement by epifaunal biota (Harding and Mann, 1999; 2005). In studies conducted in the Rio de la Plata Estuary almost all R. venosa specimens presented epibionts all over the shell (Giberto et al., 2006), which suggests an exposed lifestyle. Similar results were found by Savini et al. (2004 in Giberto et al., 2006) for specimens living on hard rock. Undetermined bryozoans (63%) and barnacles (Balanus sp., 78%) were the most frequent epibionts colonizing the shell of R. venosa; hydrozoan branches (30%) were also found (Giberto et al., 2006). Undetermined small sea anemones, chitons and polychaete tubes (Serpulidae) were found occasionally, and Polydora sp. infestations were found in some specimens (Giberto et al., 2006).
The presence of Polydora ciliata in the shells of R. venosa from Romanian waters of the Black Sea has been reported by Gutu and Marinescu (1979) but there is no evidence that this worm can compromise the fitness of this species in that area (ICES, 2004). In Chesapeake Bay, there have been reports of frequent specimens found with boring of the shell in the apical region, that corresponds to internal mud blisters characteristic of Polydora websteri infestations (ICES, 2004). These marks were clearly restricted to the early life spans of the individuals (ICES, 2004), and there is no evidence that these infestations have a detrimental effect on the survival of R. venosa.
R. venosa is a subtidal species that favours compact sandy bottoms, in which it burrows almost completely with only the siphon sticking out (Zenetos et al., 2004). Evidence from the Chesapeake Bay population suggests that this species occupies shallow hard-substrate habitats until reaching shell lengths in excess of 70 mm and then migrates into deeper habitats with sand or mud substrates where it forages on infaunal bivalves (ICES, 2004). In the Black Sea it occurs on sandy and hard bottom substrates to 40 m depth (ICES, 2004).
Wu (1988) reports that in its native range, R. venosa can exploit warm summer temperatures and avoid possible surface freezing in winter by migrating into deeper water in these regions (Mann and Harding, 2003). In the Black Sea, temperatures have a winter minima of ~7°C and a summer maxima of ~24°C. R. venosa occupies a salinity range of 25-32 ppt (Golikov, 1967), but in the Sea of Azov it can persist in areas of mean annual salinity <12 ppt (ICES, 2004). Percentage survival of R. venosa larvae is significantly less at 7 ppt that any other salinity and in its native Korean range it demonstrates large annual temperature tolerances (from 4-27°C).
ClimateTop of page
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
Means of Movement and DispersalTop of page
The main vector of long distance dispersal of the gastropod R. venosa is presumed to be shipping, and especially ballast water (ICES, 2004). Its occurrence on the mid-Atlantic coast of the USA (Chesapeake Bay) and in the Rio de la Plata Estuary was probably facilitated by the transport of larval stages in ballast water (Mann and Harding, 2003). In the North Sea, specimens have been found in the vicinity of areas with heavy shipping (Kerckhof et al., 2006).
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Economic/livelihood||Positive and negative|
Economic ImpactTop of page
R. venosa is an active predator of epifaunal bivalves and its proliferation is a serious limitation to natural and cultivated populations of oysters and mussels (Zenetos et al., 2004). It is internationally considered a serious menace to bivalve fisheries, being preferentially acclimated in estuarine/brackish water of coastal regions, where intensive bivalve harvesting usually takes place (Savini et al., 2007). In the North Sea, the regional industries for edible bivalves such as mussels Mytilus edulis, Pacific oysters Crassostrea gigas and cockles Cerastoderma edule may be at risk (Kerckhof et al., 2006; ISSG, 2007). In Chesapeake Bay, the successful local recruitment of R.venosa has raised concerns about the co-location of the invasion with a native hard clam (Mercenaria mercenaria) population that supports a local fishery worth in excess of US $3 million per year (ICES, 2004). Finally, squid fishermen in the Adriatic are particularly disturbed by the presence of this gastropod which utilises nets as spawning substratum by crawling inside, occupying all the room available and adding extra load to the draught (ISSG, 2007).
The possible effects of R. venosa in the North Sea remain uncertain, but if established, this invasive species could become a severe competitor for the native whelk Buccinum undatum. As R. venosa is known as a predator on bivalves, an impact on local aquaculture activities (on e.g. blue mussels and oysters) is also possible (Minchin et al., 2013).
Environmental ImpactTop of page
R. venosa has become established in the Black Sea with significant damage to native benthos (e.g. bivalves; notably Ostrea edulis, Pecten ponticus, and Mytilus galloprovincialis) (Mann and Harding, 2003). It has occupied an empty ecological niche exerting a significant predatory pressure on the indigenous malacofauna. Impact on bivalve populations is variable ranging from rather mild along the Romanian coast, moderate in Bulgarian and Turkish Black Sea, and severe along Russian and Ukrainian coasts where this species has been blamed for local extermination/major decline of a number of bivalves (BSEPR, 2007). Illegal bottom trawling for harvesting of R. venosa along the Black Sea shelf has raised ecological concerns with respect to the benthic communities and especially the mussel beds (Knudsen et al., 2010; Ulman et al., 2013).
In Chesapeake Bay, R. venosa shells have been found to provide shelter for the expanded range of the striped hermit crab, Clibanarius vittatus, which has demonstrated an ability to eat significant numbers of oyster spat when they reach sizes commensurate with their newly found ‘import size’ shelters (ICES, 2004). In the same area, the combination of pelagic larval dispersal and broad salinity tolerance of R. venosa potentially complicates the ability of the native oyster drill Urosalpinx cinerea to re-establish its formal range within the bay, which was lost by the events of Hurricane Agnes in 1972 (Mann and Harding, 2003). U. cinerea, unlike R. venosa has no pelagic larval stage and the introduction of the latter might inhibit this reestablishment process through competition (ICES, 2004).
The combination of broad dietary capabilities with broad salinity tolerances suggests that no substantial extant bivalve resources (oysters Crassostrea virginica, mussels Mytilus edulis, and soft shelled clams Mya arenaria) are in a spatial refuge from predation (Mann and Harding, 2003; ICES, 2004). The extremely fast growing rates of R. venosa in the Chesapeake Bay, combined with cryptic colouration, nocturnal habits, and preference for oysters as both food and habitat, offer serious cause for concern, particularly in light of ongoing oyster restoration efforts in the area (ICES, 2004). Finally, for this area the projected establishment range of R. venosa suggests continued predation pressure on Mercenaria mercenaria (ICES, 2004).
In the North Sea, the possible establishment of R. venosa would exert severe competition pressure to the native whelk Buccinum undatum, a species already suffering from organotin water pollution and heavy fishing pressure (Kerckhof et al., 2006).
In the Rio de la Plata Estuary, the organisms that have been found to encrust the shells of this gastropod are not usually found in the muddy bottoms of the estuary (Giberto et al., 2004), with the exception of barnacles colonizing small specimens of Mactra isabelleana. This reflects the lack of hard bottoms in the estuary and the importance of R. venosa shells as suitable settlement substrates for epibionts larvae (Giberto et al., 2006).
Due to the negative impact on ecosystem functioning, R. venosa has been classified as one of the most impacting alien species on ecosystem services and biodiversity (Katsanevakis et al., 2014).
Social ImpactTop of page
In the Black Sea demersal fisheries have expanded as the target has changed from fish stocks such as the almost depleted turbot (Scophthalmus maximus), mullet (Muglidiae) and whiting (Merlangius merlangus) to the invasive sea snail R. venosa. This is a unique case of a fisheries expansion where most are declining, and has led to concurrent social changes (e.g. influx of new fishers to the sector) in fishing communities such as Samsun on the Turkish Black Sea coast (BSEPR, 2007).
In the Turkish coast of the Black Sea, Aydin et al. (2016) reported that there are 207 vessels legally operating for whelk fisheries, 108 of which are using the hookah system. They found that the average CPUE was 1050 kg/day and income rate was 79%, suggesting that whelk fishing had a high economic gain. Each of the hookah vessels provided employment to an average of 3 individuals (Aydin et al., 2016).
Risk and Impact FactorsTop of page Invasiveness
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Capable of securing and ingesting a wide range of food
- Long lived
- Fast growing
- Has high reproductive potential
- Modification of natural benthic communities
- Negatively impacts aquaculture/fisheries
- Reduced native biodiversity
- Threat to/ loss of endangered species
- Threat to/ loss of native species
- Rapid growth
- Highly likely to be transported internationally accidentally
UsesTop of page
This is a species of economic value. In Japan, it is sold as seafood on fish markets (Zenetos et al., 2004), and a large market for this species as a foodstuff exists throughout its native range.
On comparing the biochemical composition of some edible marine molluscs at Canakkale coasts, Turkey, R. venosa appeared to be the best for a diet with relatively high protein and low lipid among the other examined molluscs (Celik et al., 2014). There are no studies of R. venosa supplementation of human diets. However, animal experiments indicate that supplementation of R. venosa to atherogenic diets improves the lipid profiles and the antioxidant status in serum of rats (Leontowicz et al., 2015).
In Korea, R. venosa is commercially exploited on hard sand bottoms by a mesh pot fishery (ICES, 2004).
R. venosa is well established in the benthic ecosystem of the Bulgarian, Romanian and Turkish Black Sea and has become a commercially valuable living resource. Demand for its meat on the international market has enhanced commercial fisheries initially in Turkey (1980s), and then in Bulgaria (1990s), while in Romania quickly developing medium-to-large subsistence harvesting is very likely to become an export-oriented industrial fishery in the coming years. In Ukraine, R. venosa uses are limited to local subsistence fishery and souvenir manufacture/trade (BSEPR, 2007). ICES (2004) reports that “a substantial fishery exists for the species along the Bulgarian and Turkish coasts with the product being exported to the orient”. The total stock size of R. venosa along the coast of the former USSR was estimated at 10,000 tones in the period 1988-1992 (Serobaba and Chashchin, 1995; Zaitsev and Alexandrov, 1998 in ICES 2004).
R. venosa shells are marketed as tourist souvenirs.
Its use as an alternative source for nano-bioceramic production has been examined. The results of a study using cleaned sea snail samples provided from local markets in Istanbul, showed that to produce hydroxyapatite (HA) and other bioceramic phases, hot-plate stirring method is a reliable, rapid and economic method when compared to other tedious HA production methods. Moreover, sea snail shells are very good candidate materials to produce fine powders with hotplate stirring method for various tissue engineering applications (Ozyegin et al., 2012).
In Ukrainian waters, it destroyed the oyster banks in the area of the Kerch Strait and in Karkinitsky Bay. Hence, the species has a severe impact on all ecosystem services provided by mussel and oyster biogenic reefs, i.e. food provision, water purification, coastal protection, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al., 2012).
Another reference to environmental services by this species is by Liang et al. (2004 in GISD, 2005) who states that: “R. venosa manifested the most bioaccumulation capacity of Cd (cadmium). R. venosa and the short necked clam Ruditapes philippinarum were hopeful bioindicators for monitoring Cd and Ni (nickle) pollution in waters ...”
R. venosa is a high level ecological gastropod due to its high fertility, fast growth rate, tolerance to low salinity, high temperature, water pollution and oxygen deficiency. Studies in the Black Sea (Ukraine, Romania) have shown that the intense activity of antioxidant enzymes reflects the degree of contamination, which confirms the use of this species as a bioindicator in oxidative stress, in biomonitoring of metal aquatic pollution (Moncheva et al., 2011; Jitar et al., 2015).
Uses ListTop of page
- Pollution indicator
Human food and beverage
- Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)
Similarities to Other Species/ConditionsTop of page
R. venosa is one of three species of the genus Rapana in Chinese waters and it resembles the tropical Indo-Pacific species Rapana rapiformis. It is distinguished by having a higher spire with a shoulder rather than a keel and a heavier shell with the inside of the aperture being bright orange (Zenetos et al., 2004). Another indo-pacific species Rapana bezoar is distinguished by a more scaly aperture overall and a white aperture (ICES, 2004).
Prevention and ControlTop of page
As with many introductions, the probability of observing the initial introduction event is minimal (ICES, 2004). Given that the larval stage of R. venosa is the most probable candidate for introduction via ballast water vectors, any observation of adults in a receptor region will occur after a long period of time, during which, accompanying invading individuals will have dispersed and become established elsewhere (ICES, 2004).
In the Chesapeake Bay area, an extensive public education program is in effect, and a bounty exists for R. venosa individuals delivered to the Virginia Institute of Marine Science. There is also guidance to destroy egg cases when found (ICES, 2004). The significance of public awareness concerning R. venosa invasions is evident in the ICES (2004) report on R. venosa, which states that “public education can and must be supported to underscore the potential damaging effects of this species on native species of commercial and/or ecological importance”.
Preventing the spread of marine invaders such as R. venosa that are transported by ballast water could involve the sampling and treating of ballast water systems, while knowing the potential spread of a marine invader may highlight areas at risk of invasion and indicate appropriate areas to prioritise in terms of preventing its introduction into new locations (ISSG, 2007).
Prospects for control or eradication of an invading population are bleak when physical conditions and potential prey concentrations are amenable to establishment (ICES, 2004). Attempts to target this species for control or eradication have notable weaknesses. The ICES (2004) report on R. venosa is very comprehensive and precise in this matter: “Egg case mats, although visible and often concentrated, may be spread over vast areas, and, given the large number of developing embryos per case, represent considerable propagule pressure when present even in small numbers. Larval forms are too dispersed to be considered tractable targets when free swimming in receptor environments. Identification and collection of post-settlement forms on hard substrates is difficult in complex community structures, given the probability of confusion with other gastropods is high, while total community destruction is untenable. While large epifaunal individuals are identified with comparative ease, their selective collection represents an enormous investment of diver time. Collection of infaunal individuals is tractable with commercial dredges or pots/traps designed for target infauna. Extensive dredging for disparate populations of the invader would precipitate unacceptable levels of environmental destruction with accompanying debilitation of native species. In summary, there are no proven methods currently available for control or eradication of this species should it become established in a receptor environment”.
A possible control but not eradication option is also described in the ICES (2004) report: “Investigations are in progress to examine the use of such techniques as side-scan sonar to identify significant mating aggregations on open sand substrate in the lower Chesapeake Bay. These may prove useful in guiding removal of those aggregations with reduction of propagule pressure, but they are not universally applicable and will not result in complete removal of reproductively capable or active individuals”. The probable habitat overlap between juvenile blue crabs and R. venosa in Chesapeake Bay and the predation by blue crabs on epifaunal R. venosa is a form of natural biological control which may be occurring in the area (Harding and Mann, 2003). Mud crabs and spider crabs (Libinia emarginata) also consume R. venosa. However the deliberate distribution of crabs into estuarine habitats occupied by R. venosa with no prior host range testing is unadvisable (ISSG, 2007).
The probable habitat overlap between juvenile blue crabs and R. venosa in Chesapeake Bay and the predation by blue crabs on epifaunal R. venosa is a form of natural biological control which may be occurring in the area (Harding and Mann, 2003). Mud crabs and spider crabs (Libinia emarginata) also consume R. venosa. However the deliberate distribution of crabs into estuarine habitats occupied by R. venosa with no prior host range testing is unadvisable (ISSG, 2007).
No widely effective control options to eliminate the species are available at the present time, and commercial fisheries represent the most significant way for the collection of R. venosa in numbers to implement control (ICES, 2004).
There are no obvious control measures in place to prevent continuing range expansion of R. venosa westward in the Mediterranean (ICES, 2004), although an eradication program using nets and dredges as well as a public education program were reported to be underway at the time of the publication of the ICES report (2004) on the coast of Brittany in France.
Gaps in Knowledge/Research NeedsTop of page
In the comprehensive ICES report about R. venosa (2004) the lack of specific information in many of the aspects of the biology of this species, and its significance was pointed out: “no studies of the current populations in the native range were found, no data could be found on the growth rate or longevity of R. venosa in Korea, lack of growth data on R. venosa compromises the ability to estimate the date of introduction” (ICES, 2004). These points represent only some of the missing information about this species. Mann and Harding (2003) state that “for adults of this species neither salinity tolerance nor distribution in estuarine systems of graded salinity, are well described in the literature for native or invading populations”.
ReferencesTop of page
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ContributorsTop of page
14/02/16 Updated by:
Argyro Zenetos, Institute of Marine Biological Resources & Inland Waters, Hellenic Centre for Marine Research, P.O. BOX 712, Anavissos 19013, Greece
11/12/07 Original text by:
Argyro Zenetos, Institute of Oceanography, Hellenic Centre for Marine Research, P.O. BOX 712, Anavissos 19013, Greece
Distribution MapsTop of page
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