Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Rapana venosa
(veined rapana whelk)



Rapana venosa (veined rapana whelk)


  • Last modified
  • 13 February 2020
  • Datasheet Type(s)
  • Invasive Species
  • Natural Enemy
  • Preferred Scientific Name
  • Rapana venosa
  • Preferred Common Name
  • veined rapana whelk
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Mollusca
  •       Class: Gastropoda
  •         Subclass: Caenogastropoda
  • Summary of Invasiveness
  • 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 i...

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Rapana venosa (veined rapana whelk); typical shell. Length 10cm. (a) dorsal. (b) lateral (right side). (c) ventral. (d) back. (e) front view. Samples originating from the Black Sea, nr. Sevastopol, Ukraine; ex coll. George Chernilevsky.
TitleTypical shell
CaptionRapana venosa (veined rapana whelk); typical shell. Length 10cm. (a) dorsal. (b) lateral (right side). (c) ventral. (d) back. (e) front view. Samples originating from the Black Sea, nr. Sevastopol, Ukraine; ex coll. George Chernilevsky.
Copyright©H. Zell/via wikipedia - CC BY-SA 3.0
Rapana venosa (veined rapana whelk); typical shell. Length 10cm. (a) dorsal. (b) lateral (right side). (c) ventral. (d) back. (e) front view. Samples originating from the Black Sea, nr. Sevastopol, Ukraine; ex coll. George Chernilevsky.
Typical shellRapana venosa (veined rapana whelk); typical shell. Length 10cm. (a) dorsal. (b) lateral (right side). (c) ventral. (d) back. (e) front view. Samples originating from the Black Sea, nr. Sevastopol, Ukraine; ex coll. George Chernilevsky.©H. Zell/via wikipedia - CC BY-SA 3.0
Rapana venosa (veined rapana whelk); live specimen, ventral view showing closed operculum. Black Sea. 2008.
TitleLive specimen
CaptionRapana venosa (veined rapana whelk); live specimen, ventral view showing closed operculum. Black Sea. 2008.
CopyrightPublic Domain - Released by George Chernilevsky - CC 0
Rapana venosa (veined rapana whelk); live specimen, ventral view showing closed operculum. Black Sea. 2008.
Live specimenRapana venosa (veined rapana whelk); live specimen, ventral view showing closed operculum. Black Sea. 2008.Public Domain - Released by George Chernilevsky - CC 0
Rapana venosa (veined rapana whelk); dorsal view of shell, showing general appearance.
CaptionRapana venosa (veined rapana whelk); dorsal view of shell, showing general appearance.
Copyright©Argyro Zenetos
Rapana venosa (veined rapana whelk); dorsal view of shell, showing general appearance.
ShellRapana venosa (veined rapana whelk); dorsal view of shell, showing general appearance.©Argyro Zenetos


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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 Invasiveness

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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).

R. venosa was first recorded outside its native distribution in the Black Sea in 1946. Its establishment in the Black Sea appeared to be facilitated by the general lack of competition from other predatory gastropods and an abundance of potential prey species (ICES, 2004).

In the past decades, its biogeographical range has extended towards Europe and America due to shipping (ballast water and aquaculture transfer) (Savini et al., 2007). Furthermore, its cryptic nature contributes to the improbability of observing individuals until they are large and imposing members of the benthic community (ICES, 2004).

Taxonomic Tree

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  • 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 Nomenclature

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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).

Four species of the genus Rapana are found in the literature. The veined rapa whelk R. venosa (Valenciennes, 1846), the turnip shell Rapana rapiformis (Born, 1778), the bezoar rapa whelk Rapana bezoar(Linnaeus, 1758), and the indo-Pacific species Rapana bulbosa(Solander, 1817) (Glayzer et al., 1984). No known hybrids or varieties of R. venosa exist in the literature. This species has also been described with the junior synonyms Rapana thomasiana Crosse 1861, Rapana thomasiana thomasiana (Thomas’ Rapana venosa) (ICES, 2004), Rapana pontica Nordsieck, 1969 (Zenetos et al., 2004).



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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.

Older specimens can be eroded, but typically the colour of the shell is variable from dull grey, to orange-brown, and atypically blonde, with more or less conspicuous dark brown dashes on the spiral ribs which tend to make an interrupted vein-like pattern on the cords throughout the entire shell. The aperture and columella vary from deep orange to yellow, or off-white. Additionally, spiral, vein-like colouration, varying from black to dark blue, occasionally occurs internally, originating at the individual teeth at the outer lip of the aperture. Local variation may occur in morphometry and colouration depending on substrate. Adult specimens from the Far East measure up to 180 mm in length, whereas Mediterranean and Black Sea specimens are usually less than 120 mm in length (Zenetos et al., 2004). Individuals of more than 170 mm have been collected from Chesapeake Bay, USA (ICES, 2004).

Juveniles of R. venosa may have the interior of the aperture more distinctly fluted, are more brownish, and have dark streaks along the furrows (Zenetos et al., 2004). A detailed identification and description of veliger larva development of R. venosa is found in Saglam and Duzgunes (2007).


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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 Table

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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.

Last updated: 13 Feb 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes


ChinaPresentPresent based on regional distribution.
-FujianPresent, WidespreadNative
-HebeiPresent, WidespreadNative
-JiangsuPresent, WidespreadNative
-LiaoningPresent, WidespreadNative
-ShandongPresent, WidespreadNative
-TianjinPresent, WidespreadNative
-ZhejiangPresent, WidespreadNative
JapanPresentPresent based on regional distribution.
-HokkaidoPresent, WidespreadNative
-HonshuPresent, WidespreadNative
-KyushuPresent, WidespreadNative
-Ryukyu IslandsPresent, WidespreadNative
-ShikokuPresent, WidespreadNative
TaiwanPresent, WidespreadNative
TurkeyPresent, LocalizedIntroduced1960Invasive


BulgariaPresent, WidespreadIntroduced1956Invasive
FrancePresentIntroduced1998Bay of Quiberon
GreecePresentIntroduced1986Thermaikos Gulf
ItalyPresent, WidespreadIntroduced1973Northern Adriatic; (Ionian: Crocetta and Soppelsa, 2006)
NetherlandsPresent, Few occurrencesIntroduced2005Off Scheveningen, accidental
RomaniaPresent, WidespreadIntroduced1961InvasiveBlack Sea coast
RussiaPresentPresent based on regional distribution.
-Russian Far EastPresent, WidespreadNativePeter the Great Bay
-Southern RussiaPresent, WidespreadIntroduced1947InvasiveBlack Sea coast
UkrainePresent, WidespreadIntroduced1954InvasiveBlack Sea coast
United KingdomPresentIntroduced200530 km South of the Dogger Bank, record unconfirmed (1991); established (2005)

North America

United StatesPresentPresent based on regional distribution.
-VirginiaPresent, LocalizedIntroduced1998InvasiveChesapeake Bay

Sea Areas

Atlantic - NortheastPresent, LocalizedIntroducedBay of Quiberon, France
Atlantic - NorthwestPresent, LocalizedIntroducedInvasiveChesapeake Bay. VA
Atlantic - SouthwestPresent, LocalizedIntroducedRio de la Plata estuary
Mediterranean and Black SeaPresent, WidespreadIntroducedInvasive
Pacific - NorthwestPresent, WidespreadNative
Pacific - Western CentralPresent, WidespreadNative

South America

ArgentinaPresent, LocalizedIntroduced1998Rio de la plata Estuary
BrazilPresent, LocalizedIntroducedPresent, based on regional distribution
-Rio Grande do SulPresent, LocalizedIntroduced2017
UruguayPresent, LocalizedIntroduced1998Rio de la plata Estuary

History of Introduction and Spread

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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).

In the Mediterranean Sea it was first recorded in 1973 in the northern Adriatic (Ghisotti, 1974), and Slovenia (De Min and Vio, 1997) while a separate introduction presumably via the Dardanelles Straits is held responsible for the presence of the species in the Greek northern Aegean Sea (1986) (Koutsoubas and Voultsiadou-Koukoura, 1991) and Turkish Aegean (Engl, 1995). In 2002 it was also reported from Mikhmoret Beach in Israel (Mienis, 2004).

The first confirmed sighting of R. venosa along the Brittany coast of France was in 1998 in a subtidal area in the Bay of Quiberron, although the date of first introduction remains unknown (Goulletquer, 2002). It was apparently introduced into the area as ballast in clam culture bags of Ruditapes philipinarum that were transferred from the Adriatic (Goulletquer, 2002).

The first record of R. venosa in Chesapeake Bay, USA, was a single specimen collected in June 1998, although estimates of age given by experienced malacologists, together with extensive anecdotal information from local commercial fishermen, indicate an introduction date as early as 1988 (ICES, 2004). Current population demographics in the area suggest a single introduction event (ICES, 2004).

Limited comparisons of both mitochondrial DNA, and nuclear DNA from R. venosa populations in Korea, Mediterranean Turkey and the Chesapeake Bay (Gensler et al., 2001 in ICES, 2004) shows strong evidence for a significant founder effect in the Black Sea population compared to the native range, whereas the Chesapeake Bay population appears to exhibit strong similarity to the Black Sea population (ICES, 2004). This supports the hypothesis that the species came in with ballast water from the Mediterranean (ICES, 2004).

A single female and egg capsules were found in the north of the Bahia Samborombon on Rio de la Plata (Argentina) in 1998 (ICES, 2004; Giberto et al., 2006). R. venosa successfully spread over the muddy bottoms of the estuary off Samborombon Bay and along the Uruguayan coast (Scarabino et al., 1999; Pastorino et al., 2000; Rodrigues Capitulo et al., 2002). Shipping is the mechanism suggested for the introduction of R. venosa in the Rio de la Plata Estuary (Pastorino et al., 2000;Giberto et al., 2006).



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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Albania Italy 2011 No No Ruci et al. (2014)
Argentina 1998 No No Giberto et al. (2004)
Belgium 2006 No No Kerckhof et al. (2006)
Bulgaria Japan 1954 No No Kaneva-Abadjieva (1958)
France Italy 1998 No No Goulletquer et al. (2002)
Greece 1986 No No Koutsoubas and Voultsiadou-Koukoura (1991) Introduced from the Black Sea
Israel 2002 No No Mienis (2004)
Italy 1973 No No Ghisotti (1974)
Romania Japan 1961 No No Grossu (1964)
Russian Federation Japan 1947 No No Drapkin (1963)
Slovenia Italy 1997 No No Min and Vio (1997)
Turkey 1960 No No Fischer-Piette (1960) Introduced from the Black Sea
UK 2005 No No Kerckhof et al. (2006)
Ukraine Japan 1954 No No Chukhchin (1961)
Uruguay 1998 No No Giberto et al. (2004)
USA 1988-1998 No No Harding and Mann (1999)

Risk of Introduction

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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).


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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.

Positive economic effects from R. venosa fishery are counteracted by negative ecological side-effects of destructive fishing practices used in Turkey and Bulgaria where R. venosa is fished with dredges and beam trawls, in the latter country illegally. In contrast, in Romania R. venosa is selectively fished by SCUBA divers, a sustainable method which does not disturb the habitat or involve by-catches of other animals. However, signs of over-harvesting are already evident in some areas. A new, sustainable method of harvesting Rapana is currently being trialled in Turkey, with promising results. This uses baited traps, analagous to lobster/crab pots, which offer no harm to benthic habitats, with minimal by-catches and greater control over the size/age of Rapana caught.

Habitat List

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LittoralCoastal areas Principal habitat Natural
BrackishEstuaries Secondary/tolerated habitat Harmful (pest or invasive)
BrackishLagoons Principal habitat
MarineInshore marine Principal habitat Natural
MarineBenthic zone Secondary/tolerated habitat Natural

Biology and Ecology

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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).

Reproductive Biology

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.

Environmental Requirements

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).


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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 Dispersal

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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).

Although the long planktonic larval stage of this species allows accidental transfer throughout the oceans in ships’ ballast water, accidental introduction of egg cases in hull fouling or with aquaculture products is very plausible (Kerckhof et al., 2006).

Once a founder population is established within a basin or water body, expansion of the range within the basin or propagation of the invasion front may be the result of larval dispersal from nursery areas via tidal currents (Mann et al., 2006). The potential for long range distance dispersal within a single generation remains to be determined, although recent collections of small <75 mm in length) adults on the Virginia Bay shore of the Delaware-Maryland-Virginia Peninsula suggest that a distance of tens of kilometers per generation is possible (Mann and Harding, 2003).

Observations of R. venosa massively bio-fouling immature green sea turtles Chelonia mydas confirm biotic transfer. From November 2004 to July 2011, 33 green turtles with rapa whelks attached to their carapaces were examined in the Río de la Plata Estuary, Uruguay (Lezama et al., 2013; Spotorno-Oliveira et al., 2020).

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aquaculture stock Yes Yes ICES (2004)
Host and vector organismsFouling Chelonia mydas Yes Lezama et al. (2013)
Live seafood Yes Yes ICES (2004)
Ship ballast water and sedimentMain form of transport, larval stage Yes Yes ICES (2004)

Impact Summary

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Cultural/amenity None
Economic/livelihood Positive and negative
Environment (generally) Negative
Human health None

Economic Impact

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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 Impact

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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 coloration, 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).

Rapana venosa may affect the bouyancy of the Red List endangered green turtle (Chelonia mydas) by adhering to the carapace (Spotorno-Oliveira et al., 2020).

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).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Chelonia mydas (green sea turtle)EN (IUCN red list: Endangered); USA ESA listing as endangered speciesBrazilFoulingSpotorno-Oliveira et al. (2020)

Social Impact

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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 Factors

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  • 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
Impact outcomes
  • Conflict
  • Modification of natural benthic communities
  • Negatively impacts aquaculture/fisheries
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Fouling
  • Predation
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally


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Economic Value

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).

Environmental Services

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 List

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  • Pollution indicator


  • Souvenirs

Human food and beverage

  • Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)

Similarities to Other Species/Conditions

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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).

In the Atlantic coast of the USA, found on Cape Cod and southwards, the knobbed whelk Busycon carica and the channelled whelk Busycon canliculatum are the only larger whelks that would likely be confused with R. venosa. Neither of these species have a broad columnella nor the black vein-like pattern on the shell, characteristic of R. venosa.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.


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).

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 Needs

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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”.

The current demographic data of native R. venosa populations in Asia is simply not accessible, while data on genetics and physiology are scarce. Many basic biological questions about the reproductive seasonality, egg capsules and larval development of R. venosa have still to be answered. Such biological information would be necessary for the development of techniques for re-stocking natural populations in areas where fisheries of this species exist (Saglam and Duzgunes, 2007).


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Distribution References

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Fischer-Piette E, 1960. Rapana bezoar I. in the Black Sea coast of Turkey. (Rapana bezoar I. sur la cote Turquie de la mer Noire). Hydrobiologie, Serie B5. 51.

Ghisotti F, 1974. [English title not available]. (Rapana venosa (Valenciennes), nuova ospite adriatica?). Conchilie. 10 (5-6), 125-126.

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Goulletquer P, Bachelet G, Sauriau Noel P G P, 2002. Open Atlantic coasts of Europe - a century of introduced species into French waters. In: Invasive Aquatic Species in Europe. Distribution, Impacts and Management. [ed. by Leppakoski E]. Dordrecht, Netherlands: Kluwer Academic Publishers. 276-290.

Grossu A, 1964. [English title not available]. (Die Gastropodenfauna aus der nördlichen Kleinen Walachei (Südkarpaten) und ihre biogeographischen Eigenschaften). In: Zoologische Abhandlungen - Abhandlungen und Berichte aus dem Staatlichen Museum für Tierkunde in Dresden, 14.

ICES, 2004. Alien Species Alert: Rapana venosa (veined whelk). In: Alien Species Alert: Rapana venosa (veined whelk), [ed. by Mann R, Occhipinti A, Harding M J].

Kaneva-Abadjieva V, 1958. A new harmful snail along the Bulgarian Black Sea coast. Priroda. 89-91.

Kerckhof F, Vink R J, Nieweg D C, Post J J N, 2006. The veined whelk Rapana venosa has reached the North Sea. Aquatic Invasions. 1 (1), 35-37. DOI:10.3391/ai.2006.1.1.8

Koutsoubas D, Voultsiadou-Koukoura E, 1991. The occurrence of Rapana venosa (Valenciennes, 1846) (Gastropoda: Thaididae) in the Aegean Sea. Bolletino Malacologico. 201-204.

Mienis H K, 2004. New data concerning the presence of lessepsian and indo-pacific migrants among the mollusks in the Mediterranean Sea with emphasis on the situation in Israel. In: Proceedings of the First National Malacology Conference, Izmir [Proceedings of the First National Malacology Conference, Izmir], 117-131.

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Rolán E, Bañón R, 2007. (Primer hallazgo de la especie invasora Rapana venosa y nueva información sobre Hexaplex trunculus (Gastropoda, Muricidae) en Galicia). In: Noticiario de la Sociedad Española de Malacología, 47 57-59.

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Links to Websites

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AquaNIS system on aquatic non-indigenous and cryptogenic species
EASIN (European Alien Species Information Network) developed by the European Commission’s Joint Research Centre which enables access to data on Alien Species reported in Europe.
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS) source for updated system data added to species habitat list.


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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

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