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information on the biology of species and the ecology of habitats found around the coasts and seas of the British Isles

Fan mussel (Atrina fragilis)

NBN Interactive05-09-2007

Map accurate at time of writing. Visit NBN or OBIS to view current distribution

Researched byDr Harvey Tyler-Walters Refereed byDr Dan Minchin
Authority(Pennant, 1777)
Other common names- SynonymsAtrina pectinata (Linnaeus, 1767), Pinna fragilis (Pennant, 1777), Pinna pectinata (Linnaeus, 1767)

Summary

Description

One of Britain’s largest and most threatened molluscs, growing 30-48 cm in length. The fragile shell is triangular, thin and brittle, and is tapered to a point. The shell is light yellow-brown to dark brown in colour. Fan mussels live with their pointed end buried in sediment, attached by many fine byssal threads, and the broad end protruding from the surface. Often solitary but populations occur as small groups or patches of individuals forming small beds.

Recorded distribution in Britain and Ireland

Predominantly southern and western shores of the UK from North Scotland down to Mauritania, including the Iberian Peninsula, the Mediterranean and the Channel Isles (see distribution- additional information). Recorded at Milford Haven.

Global distribution

Northern UK to Mauritania.

Habitat

Lives embedded in lower intertidal and subtidal muds, sandy muds or gravels.

Depth range

LWS to 400m

Identifying features

  • Large (30-48 cm in length by half that wide), triangular, thin and fragile shell tapering to a point.
  • Shell with conspicuous concentric growth and disturbance lines with 8-12 low, smooth, wavy ridges radiating from the beaks (umbones).
  • Occasional fluted spines.
  • Shell colour light yellow-brown to darker brown or chestnut in colour.
  • Inside of shell glossy (partly nacreous) with very dissimilar muscle scars, largest scar in middle of shell valve.
  • Valves gape at posterior (broad) end.
  • Byssus fine and abundant.
  • Shell margin smooth but often fragmented at posterior end.
  • Lives with pointed anterior end embedded in muds, sandy mud or gravel, attached to small stones or shell fragments by byssus with between one or two thirds of the shell below the sediment surface.

Additional information

According to Montagu (1803), the fan mussel has been known for its silky byssus from which expensive gloves and stockings were once made. Montagu (1803) also states that shells were, as they are now, historically brought up from trawls off the Eddystone, Plymouth, and also taken by fishermen from Torbay and on the Dorset coast.

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Biology review

Taxonomy

PhylumMolluscaSnails, slugs, mussels, cockles, clams & squid
ClassBivalviaClams, cockles, mussels, oysters, and scallops
OrderPterioidaFan mussels, hammer oysters, wing oysters and pearl oysters
FamilyPinnidae
GenusAtrina
Authority(Pennant, 1777)
Recent SynonymsAtrina pectinata (Linnaeus, 1767)Pinna fragilis (Pennant, 1777)Pinna pectinata (Linnaeus, 1767)

Biology

Typical abundanceVery low density
Male size rangeMale size at maturity
Female size rangeMedium-large(21-50cm)Female size at maturity
Growth formBivalvedGrowth ratec. 3-4cm/year
Body flexibilityNone (less than 10 degrees)Mobility
Characteristic feeding methodActive suspension feeder
Diet/food source
Typically feeds onPhytoplankton
SociabilityGregarious Environmental positionInfaunal
DependencyIndependent.
SupportsHost

Commensal Pinnotheres pinnotheres (pea crab) and Capulus ungaricus (Hungarian hat shell).

Is the species harmful?See additional information

This species may become toxic due to the accumulation of toxins during blooms of toxic algae.

Biology information

Anon (1999c) suggested that growth is relatively slow (c. 3 - 4 cm/year), based on annular growth rings in specimens from Valentia Bay, Ireland. Bulter et al. (1993) note that the growth of Pinna bicolor is indeterminate, rapid when small and slow after 2 years of age. Shell margin is easily damaged to produce sharp edges. These edges may cut the feet of bathers (Anon 1999c). Fan mussels rapidly repair shell damage (Yonge, 1953).

Community interactions
Fan mussel communities have been poorly studied in the UK and information is only available from the Mediterranean, South Australia and the USA.

Mollusc shells are often important settlement substrates for sessile organisms, or shelters/nest sites for mobile benthic animals in marine soft sediments (Kuhlmann 1998). The resource availability of shells may affect the population dynamics of these species.

Pinna bicolor hosts a species rich epifauna (Kay & Keough, 1981; Ward & Young 1984). In St Joseph Bay, Florida shells of dead Atrina rigida, predated by the horse conch, provide shelter for crabs, fish and octopus and were used as 'nests' by blennies, clingfish and toadfish (Kuhmann, 1997).

The presence of burrowing bivalves may also alter small scale flow dynamics, as is the case with Atrina zealandica (Nikora et al., 2002). However Atrina zealandica occurs in far greater densities than UK examples of Atrina fragilis. It has been found that the distance between individuals of Atrina zealandica affects the associated benthic macrofauna (Hewitt et al., 2002). The settlement of biodeposits (faeces and pseudofaeces) from Atrina zealandica is likely to have localised effects on benthic community structure (Miller et al., 2002). The same may be true of Atrina fragilis, although these effects are likely to be reduced due to far lower densities.

Pseudofaeces from pen shells results in biodeposits that reduce the variability of nematode meiofauna in Atrina zealandica beds (Warwick et al. 1997).

Habitat preferences

Physiographic preferencesOffshore seabed, Ria / Voe, Enclosed coast / Embayment
Biological zone preferencesLower circalittoral, Lower infralittoral, Sublittoral fringe, Upper circalittoral, Upper infralittoral
Substratum / habitat preferencesCoarse clean sand, Fine clean sand, Gravel / shingle, Mixed, Mud, Muddy gravel, Muddy sand, Sandy mud
Tidal strength preferencesModerately Strong 1 to 3 knots (0.5-1.5 m/sec.), Weak < 1 knot (<0.5 m/sec.)
Wave exposure preferencesSheltered, Very sheltered
Salinity preferencesFull (30-40 psu)
Depth rangeLWS to 400m
Other preferencesNo text entered
Migration PatternNon-migratory / resident

Habital Information

The present distribution and abundance of Atrina fragilis is likely to be greatly reduced compared with the historical abundance.

Distribution
Specimens occur as solitary individuals or localised patches of small groups of individuals.

Atrina fragilis was recorded in Zostera sp. beds in the Isles of Scilly. However specimens have not been found since the Zostera sp. beds were lost (Turk 1982; Turk & Seaward 1997). This species was more common in scallop areas in the early 1900s. Presumably trawling and dredging of these formerly populated regions is the reason for the decline of this species (Minchin pers. comm.). Dredging of a Pecten maximus bed off Glengad Head, Ireland, after 1975, removed many live specimens of Atrina fragilis in scallop dredges and the population of fan mussels is thought to have been destroyed by subsequent dredging (Anon 1999c).

The species is known from weak to moderately strong currents. For example in Knightstown, Valentia Island, the population is exposed to >2 knots on spring tides (Dan Minchin pers. comm.). Although usually found at full salinities it may be exposed to reduced or variable salinities for brief periods (Dan Minchin pers. comm.).

Records for Ireland were supplied by Julia Nunn and the Centre for Environmental Data and Recording (CEDaR). Records of fan mussel shells, and records before 1950, are more widespread than shown in the above map, which only shows live records after 1950.<\p>

Life history

Adult characteristics

Reproductive type Gonochoristic (dioecious) Reproductive frequency No information
Fecundity (number of eggs) No information Generation time Insufficient information
Age at maturity Insufficient information Season Insufficient information
Life span Insufficient information

Larval characteristics

Larval/propagule type - Larval/juvenile development Planktotrophic
Duration of larval stage No information Larval dispersal potential >1000m
Larval settlement period Insufficient information

Life history information

Atrina fragilis is assumed to be long lived due to its large size. Butler et al. (1993) report that the related Mediterranean genus Pinna nobilis may live up to 18 years. Dispersion is assumed to be possible over large distances due to the pelagic life history. However, Butler et al. (1993) mention evidence for genetic population sub-division in Pinna bicolor within the Gulf of St Vincent in South Australia, which suggests that effective dispersal is lower than expected. Fertilization is external and dependant on proximity of other individuals, and factors including water movement. When populations of Pinna become very sparse, as is the case in the UK, fertilization failure is likely to be significant (Anon, 1999c; Butler et al., 1993). Recruitment is likely to be sporadic due to variable larval survival and irregular, limited dispersal. Dispersal between different patches is likely to be variable (Anon, 1999c).

Sensitivity reviewHow is sensitivity assessed?

Physical pressures

 IntoleranceRecoverabilitySensitivityEvidence/Confidence
High Very low / none Very High Moderate
Atrina fragilis lives embedded in the substratum. Removal of substratum will involve removal of this species. Although they can burrow once vertical (Yonge 1953), adults cannot dig themselves back into the sediment and will not survive being uprooted. Anon (1999c) reports the destruction of a population of Atrina fragilis off Glengad Head, Ireland after 1975 by scallop dredging.
Therefore intolerance to this factor is assessed as high. Recovery is likely to be very low (see additional information), hence sensitivity has been assessed as very high.
Intermediate Low High Low
Atrina fragilis cannot burrow upwards through sediment (Yonge 1953). However 1/3 to 1/2 of the animal can protrude above the surface which, in adults, can be up to 10 - 15 cm above the sediment surface. Therefore adult specimens may not be affected by this factor at the benchmark level. However small or juvenile specimens may be smothered. Pinnids are adapted to a sedimentary life style and exhibit a powerful exhalent current and a unique ciliated waste canal to remove sediment from the mantle cavity, as would be expected from occasional smothering due to storms (Yonge 1953). Clearance of sediment from the mantle constitutes a metabolic cost that may reduce the reproductive ability (Butler et al. 1993). However, adults are likely to cleanse themselves relatively quickly. Due to the likely lethal effects of this factor on juveniles, Atrina fragilis has been assessed as intermediately intolerant of this factor. Recovery is likely to be low (see additional information), hence sensitivity has been assessed as high.
Low Very high Very Low Moderate
Pinnids are adapted to a sedimentary lifestyle and possess a unique ciliated waste canal for the removal of sediment from the mantle cavity (Yonge 1953). However, increased siltation will require increased metabolic demand on filtration and a likely decrease in growth and reproductive capacity. Thrush et al. (1999) demonstrated a decrease in the biochemical condition in Atrina zealandica with increasing sediment load in the Mahurangi Estuary, New Zealand. Along an increasing suspended sediment gradient, long term negative effects on the biomass and growth of Atrina zelandica were observed (Ellis et al., 2002). Negative effects on condition of Atrina zelandica became apparent after only 3 days of exposure to increased suspended sediment levels, and clearance rates increased with increasing sediment loading, up to a threshold level, above which clearance rates decrease rapidly. Clearance rates of suspended sediment were lower at higher sediment concentrations (Ellis et al., 2002). It may be that Atrina zelandica found in areas with naturally high sediment loading are adapted to cope better with increases in suspended sediment than those from areas with lower background sediment concentrations. None the less, very large increases in suspended sediment are still likely to be detrimental to Atrina zelandica (Hewitt & Pilditch, 2004).
Pinna bicolor and Pinna nobilis occur in sheltered areas of low turbidity. However, juveniles settle in the boundary layer and grow rapidly to escape the high levels of sediment and it is likely that Pinnids are tolerant of suspended sediment. The absence of Pinna sp. from areas of severe sediment disturbance (Bulter et al. 1993) suggests that the populations in areas of high sediment availability will be adversely affected by increased siltation. Because adults are likely to cleanse themselves relatively quickly, intolerance of Atrina fragilis to this factor has been assessed as low. Recovery is likely to be very high, hence an overall sensitivity assessment of very low.
No information
Not relevant Not relevant Not relevant Moderate
Atrina fragilis is subtidal or only exposed at extreme low water and is unlikely to experience emersion and desiccation.
Not relevant Not relevant Not relevant Moderate
Atrina fragilis is subtidal or only exposed at extreme low water and is unlikely to experience emersion.
No information
Intermediate Low High Low
The species is known from weak to moderately strong currents, for example in Knightstown, Valentia Island the population is exposed to >2 knots on spring tides (Dan Minchin pers. comm.). Increased water flow could partly uncover adults and is likely to remove some individuals from the substratum, which would not then be able to survive to re-establish themselves. Changes in current patterns are also likely to affect larval recruitment (Anon., 1999c). Therefore intolerance of this species to an increase in water flow has been assessed as intermediate. Recovery is likely to be low (see additional information), therefore sensitivity is assessed as high.
No information
Intermediate Low High Very low
Sub-tidal species such as Atrina fragilis are likely to exhibit lower temperature tolerance than intertidal species. They are likely to be intolerant of rapid temperature change indicated in this benchmark. However, no information on temperature tolerance in Atrina fragilis was found, although it has been suggested that changes in seawater temperature are likely to affect larval recruitment pattern (Anon., 1999c).
A tropical pen shell Atrina maura was found to reach maturity more quickly at higher temperatures, taking only one month (normal maturation at lower temperatures of 20 °C takes two months). However with higher temperatures, oocytes are of poor quality than at cooler temperatures (Rodriguez-Jaramillo, 2001).
Intolerance of Atrina fragilis to temperature changes has been assessed as intermediate. Recovery is likely to be low (see additional information), therefore sensitivity is assessed as high.
No information
Low Immediate Not sensitive Moderate
It has been suggested that changes in turbidity may affect Atrina fragilis (Anon, 1999c). Pinnids are adapted to a sedimentary lifestyle and possess a unique ciliated waste canal for the removal of sediment from the mantle cavity (Yonge 1953). However, increased siltation will place an increased metabolic demand on filtration and a likely decrease in growth and reproductive capacity. Thrush et al. (1999) demonstrated a decrease in the biochemical condition in Atrina zealandica with increasing sediment load in the Mahurangi Estuary, New Zealand. Pinna bicolor and Pinna nobilis occur in sheltered areas of low turbidity. However, juveniles settle in the boundary layer and grow rapidly to escape the high levels of sediment and it is likely that Pinnids are tolerant of suspended sediment. The absence of Pinna sp. from areas of severe sediment disturbance (Bulter et al., 1993) suggests that the populations in areas of high sediment availability will be adversely affected by increased siltation. Reduction in light intensity is likely to reduce phytoplankton productivity, however, it is also likely that Pinnids feed on detritus and other suspended organic matter. Therefore intolerance has been assessed as low. Recovery on return to normal conditions is likely to be immediate. Therefore this species has been deemed not sensitive to this factor.
No information
Intermediate Low High Low
Atrina fragilis occurs in sheltered or very sheltered waters (Anon 1999c; Butler et al. 1993) and can burrow into the substratum if partly uncovered by wave action or storms (Yonge 1953). Increased water flow could partly uncover adults and is likely to remove some individuals from the substratum, which would not then be able to survive to re-establish themselves. Juveniles may removed from sediment more easily than adults. Therefore intolerance of this species to an increase in wave action has been assessed as intermediate. Recovery is likely to be low (see additional information), therefore sensitivity is assessed as high.
No information
No information Not relevant No information Not relevant
There is little known about the effects of underwater sound on marine invertebrates.
No information Not relevant No information Not relevant
Bivalve molluscs perceive light and will react to sudden shading (due to a potential predator or other source) by closing their valves. Intermittent shading will, therefore interfere with feeding in subtidal populations but is unlikely to be of relevance to circalittoral populations.
High Very High Moderate
Atrina fragilis has a fragile shell, damaged easily by anchor impact or trampling by bathers. Rapido trawling for scallops (a form of beam trawl) in the Gulf of Venice resulted in the removal of organisms from the top 2 cm of sediment and a 87% reduction in Atrina fragilis abundance in the trawl tracks. Some specimens were speared on the trawl teeth and pulled from the sediment (Hall-Spencer et al. 1999). Once removed from the sediment adults cannot dig themselves back into the sediment although they can burrow once vertical (Yonge 1953). Anon (1999c) suggested that the destruction of a population of Atrina fragilis off Glengad Head, Ireland after 1975 was caused by scallop dredging. Scallop dredging and demersal trawling have been implicated in the decline in populations of this species (Anon, 1999c). Therefore, an intolerance of high has been recorded. Recoverability is likely to be very low, therefore sensitivity has been assessed as very high.
High Very High High
Rapido trawling for scallops (a form of beam trawl) in the Gulf of Venice resulted in the removal of organisms from the top 2 cm of sediment and a 87 percent reduction in Atrina fragilis abundance in the trawl tracks. Some specimens were speared on the teeth of the Rapido trawl and pulled from the sediment (Hall-Spencer et al. 1999). Once removed from the sediment adults cannot dig themselves back into the sediment, and are unlikely to survive, although they can burrow once vertical (Yonge 1953). Anon (1999c) reports the destruction of a population of Atrina fragilis off Glengad Head, Ireland after 1975 by scallop dredging. Anon (1999c) regard pinnids to be intolerant and at threat from benthic trawling or dredging. The decline in the Mediterranean (Richardson et al. 1999) and its loss from inlets in south west England (Anon 1999c) suggests that any recovery from disturbance would be slow. Therefore intolerance has been assessed as high. Recovery has been recorded as very low (see also additional information), hence sensitivity is very high.

Chemical pressures

 IntoleranceRecoverabilitySensitivityEvidence/Confidence
No information Not relevant No information Not relevant
Insufficient
information
Heavy metal contamination
Low No information Moderate Low
Anon (1999c) suggested that Atrina fragilis may be affected by pollutants such as TBT (tri-butyl tin).
Reid & Brand (1989) describe kidney gigantism and nephroliths (calcium or iron granules) in Pinna bicolor. Their role in removing excess calcium or heavy metals and potential detoxification is unclear. Ward & Young (1983) examined changes in epifauna of Pinna bicolor due to heavy metal contamination in Spence Gulf, south Australia. They state that Pinna bicolor is tolerant of high concentrations of heavy metals in sediments near a lead smelter and contains high body loads of heavy metals. The occurrence of populations of this species in heavy metal contaminated sediment suggests that it is not sensitive. However, the body burden of Pinna bicolor was not given and no citation provided for the information. The studied population may represent a localised adaptation. Due to incomplete information, intolerance has been recorded as low, yielding a moderate sensitivity value.
Hydrocarbon contamination
No information No information No information Not relevant
Insufficient
information
Radionuclide contamination
No information No information No information Not relevant
Insufficient
information
Changes in nutrient levels
Low No information Moderate Low
Pinnids are mainly found in sheltered oligotrophic (low nutrient) waters (Butler et al. 1993), and they filter continuously, presumably an adaptation to low food availability. A small population of Atrina fragilis exists near a sewage discharge in Dingle Harbour (Dan Minchin pers comm.). An increase in nutrients is likely to increase phytoplankton production in the short term, which may benefit larvae and juveniles. Therefore intolerance has been assessed as low. Insufficient
information was found on recover from excess nutrients, therefore a moderate sensitivity value has been recorded.
No information No information Very Low Very low
Atrina fragilis occurs subtidally at full salinity, however the infralittoral populations may experience variable salinity. Dan Minchin (pers. comm.) has suggested that Atrina fragilis may be exposed to reduced or variable salinities for brief periods. It is likely, however, that this species would be intolerant of reduced salinity. A tropical pen shell Atrina maura, had been found to have a wide range of halotolerance, from 16-50 (Leyva-Valencia et al., 2001). Insufficient
information was found to make an assessment.
No information
No information No information No information Not relevant
Butler et al. (1993) state that Pinna bicolor and Pinna nobilis remain active at low oxygen concentrations and open their valves widely. Neither species stop pumping or respire anaerobically. Although bivalves are generally resistant to anoxia and can respire anaerobically no information concerning tolerance to anoxia in Pinnids was found.

Biological pressures

 IntoleranceRecoverabilitySensitivityEvidence/Confidence
Low Very high Very Low Low
The Pinnids are parasitized by the Pea crab (Pinnotheridae) (Yonge 1953) . Butler et al. (1993) state that Pinna bicolor and Pinna nobilis harbour macroscopic commensals or parasites of unknown effect, although an unidentified parasitic microbe has been recorded as causing castration of Pinna nobilis. Intolerance is therefore assessed as low, recovery very high, and sensitivity low.
No information No information No information Not relevant
Crepidula sp. may have had some impact on near shore populations of Atrina fragilis on the south coast of England (Dan Minchin pers comm.).
High Low High Moderate
In Spain, pinnids may be collected for consumption, used as bait, or for use as souvenirs. In the Bay of Naples the byssus threads were historically used for making glues. In the Pacific, declines in production have occurred as a result of exploitation in other species of penshell (Cardoza-Velasco & Maeda-Martinez, 1997) Populations in the UK are too sparse to sustain any harvesting, and are protected by a Biodiversity Action Plan and under the Wildlife & Countryside Act 1981. Based on the UK population size, intolerance has been assessed as high. Due to predicted slow growth and poor fertilisation/recruitment, recovery has been assessed as low, providing a high sensitivity assessment.
No information No information No information Not relevant
In the UK Atrina fragilis was more common in scallop beds in the early 1900s then at present. Presumably trawling and dredging of theses formerly populated regions is the reason for the decline of this species (Minchin pers. comm.). Dredging of a Pecten maximus bed off Glengad Head, Ireland, after 1975, removed many live specimens of Atrina fragilis in scallop dredges and the population of fan mussels is thought to have been destroyed by subsequent dredging (Anon 1999c).
In the Adriatic queen scallop (Aequipecten opercularis) trawl fishery, Atrina fragilis incurred more damage as a result of the fishing and sorting process than any other species of bycatch (Pranovi et al., 2001). Rapido trawling (a form of beam trawl) for scallops in the Gulf of Venice resulted in the removal of organisms from the top 2 cm of sediment and an 87% reduction in Atrina fragilis abundance in the trawl tracks. Some specimens were speared on the trawl teeth and pulled from the sediment (Hall-Spencer et al. 1999). Once removed from the sediment adults can not dig themselves back into the sediment although they can burrow once vertical (Yonge 1953). Anon (1999c) reports the destruction of a population of Atrina fragilis off Glengad Head, Ireland, by scallop dredging after 1975.
Pinnids in the Mediterranean are associated with seagrass beds, the removal of which has been linked to the decline in Pinnid populations (Richardson et al 1999). However, Atrina fragilis bed communities are little studied in the UK.

Additional information

The UK populations of %Atrina fragilis% are poorly studied and most information has been derived from studies of Mediterranean and Australasian Pinnid species.

Recovery
Penshells are vulnerable to overexploitation due to their long life, slow growth, limited reproductive output and sporadic recruitment (Butler et al., 1993). Rapid growth and rapid shell repair in Pinnids suggest high metabolic demand, that may result in reduced gamete production (Anon 1999; Butler et al., 1993), consistent with a long-lived species. Larval dispersal may be limited or irregular and larval survival variable (Butler et al., 1993). Fertilization efficiency in patchy populations of low density may also be low (Anon, 1999c). Therefore, with the exception of embayments and inlets where larvae may be trapped, recruitment of %Atrina fragilis% is poor and variable in comparison with other bivalve species (Anon, 1999c). However, surviving adults increase the possibility of fertilization and local recruitment. The decline in the Mediterranean (Richardson et al., 1999) and its loss from inlets in south west England (Anon 1999c) suggests that any recovery from disturbance would be slow. Anon (1999c) suggests that changes in factors that shorten the adult life of this species cannot be compensated for by an immediate reproductive response and recruitment.

Importance review

Policy/legislation

Wildlife & Countryside ActSchedule 5, section 9
UK Biodiversity Action Plan Priority
Species of principal importance (England)
Species of principal importance (Wales)
Priority Marine Features (Scotland)

Status

National (GB) importanceNot rare/scarceGlobal red list (IUCN) category-

Non-native

Native-
Origin- Date Arrived-

Importance information

The legendary 'cloth of gold' was made from the long fine byssus threads of pinnids (Osing, 1968, Turk, 1982). Curiously, in past times, although seldom captured, many fishermen considered Atrina fragilis unclean, and returned it directly to the sea because the byssus was so similar to human hair that it was thought that they had consumed drowned sailors (Dan Minchin, pers comm.). Mediterranean Pinna nobilis may be taken for bait, consumption and for the curio trade. The byssus of pinnids was collected in Bay of Naples for making glues (Dan Minchin pers comm.). The community structure of UK fan mussel beds is poorly studied.

Bibliography

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  7. Hall-Spencer, J.M., Froglia, C., Atkinson, R.J.A & Moore, P.G., 1999. The impact of Rapido trawling for scallops, Pecten jacobaeus (L.), on the benthos of the Gulf of Venice. ICES Journal of Marine Science, 56, 111-124
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Citation

This review can be cited as:

Tyler-Walters, H., 2007. Atrina fragilis Fan mussel. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from: http://www.marlin.ac.uk/species/detail/1157

Last Updated: 05/09/2007