Distribution data supplied by the Ocean Biodiversity Information System (OBIS). To interrogate UK data visit the NBN Atlas.Map Help
Researched by | Dr Harvey Tyler-Walters | Refereed by | Prof. R. Seed |
Authority | Linnaeus, 1758 | ||
Other common names | - | Synonyms | - |
The shell is inequilateral and roughly triangular in outline, however, shell shape varies considerably with environmental conditions. Shell smooth with a sculpturing of concentric lines but no radiating ribs. The ligament is inconspicuous. The shell colour varies, usually purple or blue but sometimes brown. Length varies, specimens usually ranging from 5 -10 cm although some populations never attain more than 2-3 cm, and the largest specimens may reach 15 -20 cm. Mytilus edulis may be confused with the Mediterranean mussel Mytilus galloprovincialis.
Mytilus edulis and Mytilus galloprovincialis often occur in the same location in the northern range of Mytilus galloprovincialis. As they both show great variation in shell shape due to environmental conditions (Seed, 1968, 1992), they are often difficult to distinguish. In addition, they may hybridize. However, in Mytilus galloprovincialis:
Note no single morphological characteristic can be used to separate Mytilus species (Gosling, 1992c; Seed, 1992, 1995). Recent evidence suggests that there are only three lineages of the genus, Mytilus edulis, Mytilus galloprovincialis and Mytilus trossulus, although some authorities suggest that all of the smooth shelled mussels belong to the same species (for discussion see Seed, 1992).
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Phylum | Mollusca | Snails, slugs, mussels, cockles, clams & squid |
Class | Bivalvia | Clams, cockles, mussels, oysters, and scallops |
Order | Mytilida | Mussels & crenellas |
Family | Mytilidae | |
Genus | Mytilus | |
Authority | Linnaeus, 1758 | |
Recent Synonyms |
Typical abundance | Moderate density | ||
Male size range | |||
Male size at maturity | |||
Female size range | Medium(11-20 cm) | ||
Female size at maturity | |||
Growth form | Bivalved | ||
Growth rate | See additional text. | ||
Body flexibility | None (less than 10 degrees) | ||
Mobility | |||
Characteristic feeding method | Active suspension feeder | ||
Diet/food source | |||
Typically feeds on | Bacteria, phytoplankton, detritus, and dissolved organic matter (DOM). | ||
Sociability | |||
Environmental position | Epilithic | ||
Dependency | Independent. | ||
Supports | Host several parasites and commensals, see additional information and Bower (1992) and Bower & McGladdery (1996) for review. | ||
Is the species harmful? | No Edible (but see 'public health' under additional information). |
Mytilus edulis is one of the most extensively studied marine organisms. Therefore, this review is based on comprehensive reviews by Gosling (ed.) (1992a), Bayne, (1976b), Newell (1989), and Holt et al. (1998). Where appropriate the original source references in these reviews are given.
Mytilus edulis is gregarious, and at high densities forms dense beds of one or more (up to 5 or 6) layers, with individuals bound together by byssus threads. Young mussels colonize spaces within the bed increasing the spatial complexity, and the bed provides numerous niches for other organisms (see importance). Overcrowding results in mortality as underlying mussels are starved or suffocated by the accumulation of silt, faeces and pseudofaeces, especially in rapidly growing populations (Richardson & Seed, 1990). Death of underlying individuals may detach the mussel bed from the substratum, leaving the bed vulnerable to tidal scour and wave action (Seed & Suchanek, 1992).
Growth rates
Growth rates in Mytilus spp. are highly variable. Part of this variation is explained by genotype and multilocus heterozygosity (Gosling, 1992b) but the majority of variation is probably environmentally determined. The following factors affect growth rates in Mytilus spp. Several factors may work together, depending on location and environmental conditions (Seed & Suchanek, 1992) or the presence of contaminants (see sensitivity, e.g. Thompson et al., 2000):
For example, in optimal conditions Mytilus edulis can grow to 60 -80mm in length within 2 years but in the high intertidal growth is significantly lower, and mussels may take 15 -20 years to reach 20 -30mm in length (Seed & Suchanek, 1992). Bayne et al. (1976) demonstrated that between 10-20 °C water temperature had little effect on scope for growth. Latitudinal variation in temperature influences shell structure in Mytilus species (Carter & Seed, 1998).
Predation and mortality
Several factors contribute to mortality and the dynamics of Mytilus edulis populations, including temperature, desiccation, storms and wave action, siltation and biodeposits, intra- and interspecific competition, and predation. But predation is the single most important source of mortality.
Many predators target specific sizes of mussels and, therefore, influence population size structure. The vulnerability of mussels decreases as they grow, since they can grow larger than their predators preferred size. Mytilus sp. may be preyed upon by neogastropods such as Nucella lapillus, starfish such as Asterias rubens, the sea urchin Strongylocentrotus droebachiensis, crabs such as Carcinus maenas and Cancer pagurus, fish such as Platichthys flesus (plaice), Pleuronectes platessa (flounder) and Limanda limanda (dab), and birds such as oystercatcher, eider, scooter, sandpiper, knot, turnstone, gulls and crows (Seed & Suchanek, 1992; Seed, 1993). Important predators are listed below.
Epifauna and epiflora
Fouling organisms, e.g. barnacles and seaweeds, may also increase mussel mortality by increasing weight and drag, resulting in an increased risk of removal by wave action and tidal scour. Fouling organisms may also restrict feeding currents and lower the fitness of individual mussels. However, Mytilus edulis is able to sweep its prehensile foot over the dorsal part of the shell (Thiesen, 1972, Seed & Suchanek, 1992). Fouling by ascidians may be a problem in rope-cultured mussels (Seed & Suchanek, 1992).
Diseases and parasites
The polychaete Polydora ciliata may burrow into the shell of Mytilus edulis, which weakens the shell leaving individuals more susceptible to predation by birds and shore crabs resulting in significant mortality, especially in mussels >6 cm (Holt et al., 1998).
Bower (1992), concluded that, although most parasites did not cause significant mortality, several species of parasite found in mussels could also infect and cause mortality in other shellfish. This suggested that mussel populations may be reservoirs of disease for other shellfish (see sensitivity or reviews by Bower, 1992; Bower & McGladdery, 1996).
Public heath
Mytilus edulis is a filter feeding organism, which collects algae, detritus and organic material for food but also filters out other contaminants in the process. Shumway (1992) noted that mussels are likely to serve as vectors for any water-borne disease or contaminant. Mussels have been reported to accumulate faecal and pathogenic bacteria and viruses, and toxins from toxic algal blooms (see Shumway, 1992 for review). Bacteria may be removed or significantly reduced by depuration (removing contaminated mussels into clean water), although outbreaks of diseases have resulted from poor depuration and viruses may not be removed by depuration. Recent improvements in waste water treatment and shellfish water quality regulations may reduce the risk of bacterial and viral contamination. Shellfish should also be thoroughly cooked, not 'quick steamed', to ensure destruction of viruses (Shumway, 1992). The accumulation of toxins from toxic algal blooms may result in paralytic shellfish poisoning (PSP), diarrhetic shellfish poisoning (DSP) or amnesic shellfish poisoning (ASP). These toxins are not destroyed by cooking. Shumway (1992) suggested that mussels should only be collected from areas routinely monitored by public health agencies, or obtained from approved sources and never harvested from waters contaminated with raw sewerage.
Physiographic preferences | Open coast, Strait / sound, Sea loch / Sea lough, Ria / Voe, Estuary, Enclosed coast / Embayment |
Biological zone preferences | Lower eulittoral, Mid eulittoral, Sublittoral fringe, Upper eulittoral, Upper infralittoral |
Substratum / habitat preferences | Artificial (man-made), Bedrock, Biogenic reef, Caves, Crevices / fissures, Large to very large boulders, Mixed, Muddy gravel, Muddy sand, Rockpools, Sandy mud, Small boulders, Under boulders |
Tidal strength preferences | Moderately Strong 1 to 3 knots (0.5-1.5 m/sec.), Strong 3 to 6 knots (1.5-3 m/sec.), Weak < 1 knot (<0.5 m/sec.) |
Wave exposure preferences | Exposed, Moderately exposed, Sheltered, Very exposed, Very sheltered |
Salinity preferences | Full (30-40 psu), Reduced (18-30 psu), Variable (18-40 psu) |
Depth range | Intertidal to ca. 5m. |
Other preferences | No text entered |
Migration Pattern | Non-migratory / resident |
Factors affecting zonation
Although sometimes abundant in the subtidal Mytilus edulis is primarily an intertidal species. Mytilus edulis can withstand extreme wave exposure, maintaining byssal attachment in high energy environments (Seed & Suchanek, 1992). The upper limit of Mytilus edulis populations on rocky shores is determined by its tolerance of temperature and desiccation, which may be synergistic, i.e. sudden mass mortalities at the upper limit of intertidal mussel beds are often associated with prolonged periods of unusually high temperatures and desiccation stress (Seed & Suchanek, 1992). Recruitment or movement into cracks, crevices or pools provides some protection from extremes of temperature and desiccation as well as from storms. Mytilus edulis is relatively tolerant of extreme cold and freezing, surviving a drop in tissue temperature to minus 10 °C (Williams, 1970). However, Bourget (1983) noted that cyclic exposures to sublethal temperatures e.g. minus 8 °C every 12.4hrs resulted in death after 3-4 cycles. This suggests that Mytilus edulis can survive occasional, sharp frost events, but may succumb to consistent very low temperatures over a few days (see sensitivity to temperature change).
Reproductive type | Gonochoristic (dioecious) | |
Reproductive frequency | Annual protracted | |
Fecundity (number of eggs) | >1,000,000 | |
Generation time | 1-2 years | |
Age at maturity | 1-2 years | |
Season | April - September | |
Life span | See additional information |
Larval/propagule type | - |
Larval/juvenile development | Planktotrophic |
Duration of larval stage | 1-6 months |
Larval dispersal potential | Greater than 10 km |
Larval settlement period | See additional information |
The MarLIN sensitivity assessment approach used below has been superseded by the MarESA (Marine Evidence-based Sensitivity Assessment) approach (see menu). The MarLIN approach was used for assessments from 1999-2010. The MarESA approach reflects the recent conservation imperatives and terminology and is used for sensitivity assessments from 2014 onwards.
Intolerance | Recoverability | Sensitivity | Evidence/Confidence | |
High | High | Moderate | High | |
Removal of the substratum, be it rock or sediment, will entail removal of the entire population and its associated community. Therefore, an intolerance of high has been recorded. Recovery may occur rapidly through good annual recruitment. However, examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. Recoverability will depend on recolonization by movement of young or juvenile Mytilus edulis from high shore or filamentous algal populations or recruitment of larvae settling directly on the new substratum. A single good recruitment event may return the population to prior levels within 1 -5 years, and an intolerance of high has been recorded. However, recoverability may be protracted in some circumstances (see additional information below). | ||||
Intermediate | High | Low | Low | |
Although apparently sedentary, Mytilus edulis is able to move some distance to change its position on the shore or within a bed or to resurface when buried by sand (Holt et al., 1998). Burial of Mytilus edulis beds by large-scale movements of sand, and resultant mortalities have been reported from Morecambe Bay, the Cumbrian Coast and Solway Firth (Holt et al., 1998). Daly & Mathieson (1977) suggested that the lower limit of Mytilus edulis populations at Bound Rock, USA, was determined by burial or abrasion by shifting sands. Dare (1976) noted that individual mussels swept or displaced from a mussel beds rarely survived, since they either became buried in sand or mud, or were scattered and eaten by oystercatchers. Dare (1976) reported that mussel beds accumulated ca. 0.4-0.75m of 'mussel mud' (a mixture of silt, faeces, and pseudo-faeces) between May and September 1968 and 1971 in Morecambe Bay. Young mussels moved upwards becoming lightly attached to each other, but many were suffocated (Dare, 1976). Therefore, it appears that mussels are able to move upwards through accumulated sediment, but that a proportion will succumb and so an intolerance of intermediate has been recorded.
Recovery may occur rapidly through good annual recruitment. However, examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. | ||||
Low | Immediate | Not sensitive | High | |
Moore (1977) reported that Mytilus edulis was relatively tolerant of turbidity and siltation, thriving in areas that would be harmful to other suspension feeders. Mytilus edulis possesses efficient shell cleaning and pseudofaeces expulsion mechanisms to remove silt (Moore, 1977), although it should be noted that pseudofaeces production involves an energetic burden (Navarro & Widdows, 1997). De Vooys (1987) examined the removal of sand from the mantle cavity and reported rapid discharge within 15min, with an exponential decrease over the next 4 hrs and slow discharge over 48 hrs. Purchon (1937) reported that Mytilus edulis died after an average of 13 days exposure to ca. 1200 mg/l suspended sediment (mud) but survived the length of the experiment (un-stated but >25 days) at 440 mg/l. Widdows et al. (1998) also noted that feeding rate was not reduced by current velocities up to 70 cm/s per se but by the resultant suspended sediment (at >50 mg/l ). Therefore, Mytilus edulis is probably of low intolerance to a change in suspended sediment at the benchmark level ( ±100 mg/l for 1 month). Recoverability is recorded as immediate given the mussels ability to discharge sand form the mantle cavity reported by De Vooys (1987). | ||||
Low | Immediate | Not sensitive | ||
A decrease in suspended sediment, especially organic particulates, could potentially reduce the food available to Mytilus edulis and hence its growth rate. Therefore, an intolerance of low has been recorded. Similarly, a recovery of immediate has been recorded. | ||||
Low | Immediate | Not sensitive | Moderate | |
The upper limit of Mytilus populations is primarily controlled by the synergistic effects of temperature and desiccation (Suchanek, 1978; Seed & Suchanek, 1992; Holt et al., 1998). For example, on extremely hot days in the summers of 1974 -1976 on Strawberry Island, Washington State, Suchanek (1978) reported mass mortality of Mytilus trossulus (as edulis) at the upper edge of the mussel bed. Mortality decreased down the shore. The upper limit of mussels fluctuated, increasing up the shore in winter and decreasing again in summer (Suchanek, 1978). British Mytilus edulis have a sustained upper thermal tolerance limit of 29 °C (Almada-Villela et al., 1982) and occur in the upper eulittoral. Holt et al. (1998) suggested that tolerance of high temperatures and desiccation explained the upper limit of Mytilus edulis on the high shore. Therefore, Mytilus edulis is likely to exhibit a relatively low intolerance to changes in desiccation at the level of the benchmark although individuals at the upper limit of the range are probably more vulnerable to desiccation. | ||||
Low | Very high | Very Low | Low | |
Mytilus edulis can only feed when immersed, therefore, changes in emergence regime will affect individuals ability to feed and their energy metabolism. Growth rates decrease with increasing shore height and tidal exposure, due to reduced time available for feeding and reduced food availability, although longevity increases (Seed & Suchanek, 1992; Holt et al., 1998). Therefore, there will be a position on the shore where the energetic cost of metabolism is not met by feeding. Baird (1966) estimated that the point of zero growth occurred at 55% emergence but this value will vary between shores depending on local conditions, e.g. wave splash (Baird, 1966; Holt et al., 1998). Increased emergence will expose mussel populations to increased risk of desiccation and increased vulnerability to extreme temperatures, potentially reducing their upper limit on the shore, and reducing their extent in the intertidal. But Mytilus edulis inhabits a wide range of shore heights and is probably relatively tolerant of changes of emergence at the benchmark level (change in emergence of 1hr for 1 year). Therefore, an intolerance of low has been reported. Similarly, once the prior emergence regime returns, the population will probably recover prior condition with a few months. | ||||
Low | Very high | Very Low | Low | |
Mytilus edulis can only feed when immersed, therefore, changes in emergence regime will affect individuals ability to feed and their energy metabolism. Growth rates decrease with increasing shore height and tidal exposure, due to reduced time available for feeding and reduced food availability, although longevity increases (Seed & Suchanek, 1992; Holt et al., 1998). Therefore, there will be a position on the shore where the energetic cost of metabolism is not met by feeding. Baird (1966) estimated that the point of zero growth occurred at 55% emergence but this value will vary between shores depending on local conditions, e.g. wave splash (Baird, 1966; Holt et al., 1998). Decreased emergence, may allow the population to colonize further up the shore but exposes the lower limit of the population to increased predation, so that the population may effectively, move up the shore. But Mytilus edulis inhabits a wide range of shore heights and is probably relatively tolerant of changes of emergence at the benchmark level. Therefore an intolerance of low has been reported. Once the prior emergence regime returns, the population will probably recover with a few months. | ||||
Low | Very high | Very Low | Moderate | |
Widdows et al. (1998) showed that Mytilus edulis beds in sheltered conditions (based on field annular flumes measurements) reduced sediment erosion. Widdows et al. (1998) also noted that feeding rate was not reduced by current velocities up to 70 cm/s per se but by the resultant suspended sediment (at >50 mg/l ). It should also be noted that mussels probably benefit from high current velocities to supply food (suspended particulates, benthic diatoms and phytoplankton). As mussel beds increase in size and depth, individual mussels become increasingly attached to each other rather than the substratum. As a result, the bed may become destabilised and susceptible to removal by wave action or tidal scour, although mussels at the edge of the beds are often more strongly attached than mussels within the bed (Seed & Suchanek, 1992). Young (1985) demonstrated that byssal thread production (and hence attachment) increased with increasing water agitation. Mussels were able to increase their byssal attachment by 25% within 8 hours of a storm commencing. Young (1985) also reported that mussels were able to withstand shock or surges of up to 16 m/s (ca 30 knots). Young (1985) concluded that mussels would be susceptible to sudden squalls and surges, which may sweep them off rocks. Therefore, storms may cause significant mortality in mussel beds (see wave exposure below). Mytilus edulis populations are found in weak to strong tidal streams, suggesting low intolerance to change in water flow rate, although, their intolerance probably owes more to the nature of the substratum than the strength of their attachment. Individuals attached to solid substrata (rock) are likely to be less intolerant than individuals attached to boulders, cobbles or sediment. Overall, Mytilus edulis can attach and grow on a variety of substrata in a variety of water flow regimes, and an intolerance of low has been reported. It should be noted that on sedimentary shores, mussel beds are probably more intolerant of increased water flow due to removal of the sediment. Once the prior water flow regime returns, the population will probably recover within a few months. | ||||
Low | Very high | Very Low | Low | |
Mytilus edulis probably benefits from high current velocities to supply food (suspended particulates, benthic diatoms and phytoplankton). A decrease in water flow is likely to decrease food availability. However, Mytilus edulis populations are found in weak to strong tidal streams, suggesting low intolerance to change in water flow rate, although, their intolerance probably owes more to the nature of the substratum than the strength of their attachment. Overall, an intolerance of low has been reported due their distribution in a variety of water flow regimes. Once the prior water flow regime returns, the population will probably recover within a few months. | ||||
Low | Very high | Very Low | High | |
In the British Isles an upper, sustained thermal tolerance limit of about 29 °C was reported for Mytilus edulis (Read & Cumming, 1967; Almada-Villla et al., 1982). But Seed & Suchanek (1992) noted that European populations were unlikely to experience temperatures greater than about 25 °C. Bayne et al. (1976) demonstrated that between 10 -20 °C water temperature had little effect on scope for growth. Mytilus edulis is generally considered to be eurythermal and an intolerance of 'low' has been recorded. Similarly, once the prior temperature regime returns, the population will probably recover any loss of condition within a few months. | ||||
Low | Very high | Very Low | High | |
Mytilus edulis can withstand extreme cold and freezing, surviving when its tissue temperature drops to -10 °C (Williams, 1970; Seed & Suchanek, 1992) or exposed to -30°C for as long as six hours twice a day (Loomis, 1995). In the laboratory, median lethal temperatures (MLT) of -16 °C after 24 hrs were estimated for large individuals (>3mm) while juveniles (<1.5mm) had an MLT of -12.5 °C. As expected, reducing the exposure time increased the MLT (Bourget, 1983). Bourget (1983) also reported that cyclic exposure to otherwise sublethal temperatures, e.g. -8 °C every 12.4 hrs resulted in significant damage and death after 3-4 cycles. This suggests that Mytilus edulis can survive occasional, sharp frost events, but may succumb to consistent very low temperatures over a few days. Mytilus edulis was relatively little affected by the severe winter of 1962/63, with 30% mortality reported from south-east coasts of England (Whitstable area) and ca. 2% from Rhosilli in south Wales (Crisp (ed.),1964). Crisp (ed.) (1964) noted that most mortality resulted from predation on individuals weakened or moribund due to the low temperatures rather than the temperature itself. Loomis (1995) reported that freezing tolerance increased after acclimation, high salinity and aerial exposure or anoxia. Freezing tolerance increased in winter, presumably due to increased isolation from the environment (shell closure), commensurate anaerobic conditions and increased mantle cavity fluid salinity with respect to the environment. Increased freezing tolerance may result from the accumulation of amino acids (e.g. taurine, glycine and alanine) involved in maintenance of osmotic balance in high salinities, and the end products of anaerobic metabolism (e.g. strombine, and octopine), which have been shown to also act as cryoprotectants (Loomis, 1995) (see oxygenation). Although Mytilus edulis may be intolerant of prolonged freezing temperatures, it is generally considered to be eurythermal and an intolerance of 'low' at the level of the benchmark is recorded. Similarly, once the prior temperature regime returns, the population will probably recover any loss of condition within a few months. | ||||
Tolerant | Not relevant | Not sensitive | Not relevant | |
Increased turbidity may reduce phytoplankton primary productivity, therefore reducing the food available to Mytilus edulis but mussels use a variety of food sources and the effects are likely to be minimal. Therefore, this species is probably tolerant to changes in turbidity. | ||||
Tolerant | Not relevant | Not sensitive | Not relevant | |
Decreased turbidity may increase phytoplankton primary productivity, therefore potentially increasing the food available to Mytilus edulis but mussels use a variety of food sources and the effects are likely to be minimal. Therefore, this species is probably tolerant to changes in turbidity. | ||||
Intermediate | High | Low | Moderate | |
Mytilus edulis populations are found in sheltered to wave exposed shores, suggesting low intolerance to change in wave exposure. Their intolerance probably owes more to the nature of the substratum than the strength of their attachment. Individuals attached to solid substrata (rock) are likely to be less intolerant than individuals attached to boulders, cobbles or sediment. Lewis (1964) noted that Mytilus edulis are favoured by damp conditions. Therefore, as wave exposure increases on rocky shores, barnacles and fucoids are replaced by mussel dominated communities. Storms and tidal surges are known to destroy mussel beds, often over hundreds of hectares in the Wash, Morecambe Bay and the Wadden Sea. Mussel beds persist in sheltered areas whereas beds in exposed areas are more dynamic (Holt et al., 1998). With increasing wave exposure mussel beds become increasingly patchy and dynamic. Young (1985) demonstrated that byssus thread production (and hence attachment) was increased by water agitation. Mussels were able to increase their byssal attachment by 25% within 8 hours of a storm commencing. Young (1985) concluded that mussels would be susceptible to sudden squalls and surges, which may sweep them off rocks. Mytilus edulis beds may also be damaged by wave driven logs or equivalent debris (Seed & Suchanek, 1992). Intense mussel settlement may lead to choking and death of underlying mussels causing the population to loosen its attachment to the substratum (Seed, 1969b). Competition for space, especially in areas of rapid growth, may lead to the formation of hummocks, in which individuals mussels may not be attached directly to the substratum. As a result, the population may become unstable and vulnerable to removal by rough seas (Seed, 1969b). Although mussel populations are found from wave exposed to sheltered shores, their intolerance to wave exposure is partly dependant on their substratum, and the size and density of the mussel bed, therefore, an intolerance of intermediate has been recorded to represent the increased susceptibility of mussel beds to damage by wave action. Recovery may occur rapidly through good annual recruitment but examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). A recoverability of 'high' has been recorded. | ||||
Intermediate | High | Low | Moderate | |
Mytilus edulis populations are found in sheltered to wave exposed shores, suggesting low intolerance to change in wave exposure. Their intolerance probably owes more to the nature of the substratum than the strength of their attachment. Lewis (1964) noted that rocky shore Mytilus edulis populations are favoured by damp conditions. Therefore, as wave exposure increases on rocky shores, barnacles and fucoids are replaced by mussel dominated communities. However, on rocky shores, as wave exposure decreases mussels are replaced by barnacle and fucoid dominated shores, possibly due to increased desiccation and predation (presumably by the dogwhelk Nucella lapillus). On wave sheltered sedimentary shores decreased wave exposure (i.e. sheltered to very sheltered) is likely to have little affect on mussel beds. Therefore, sheltered shore mussels beds are probably of low intolerance to decreased wave exposure, and may less patchy and more stable (persistent). However, on rocky shores decreased wave exposure may lead to a reduction in population density and dominance of the shore by barnacles and fucoids, therefore an overall intolerance of intermediate has been recorded. Recovery may occur rapidly through good annual recruitment but examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. | ||||
Tolerant* | Not relevant | Not sensitive* | Low | |
Mytilus edulis may detect slight vibrations in its immediate vicinity and probably detects predators by touch (on the shell) or by scent. Therefore, it is likely to be insensitive to noise disturbance at the levels of the benchmark. Birds are major predators and several species are highly intolerant of disturbance by noise. Noise at the level of the benchmark may disturb predatory birds, so that the mussel populations may benefit indirectly. | ||||
Tolerant* | Not relevant | Not sensitive* | Low | |
Mytilus edulis can probably detect changes in light commensurate with shading by predators but its visual acuity is probably very limited and it is unlikely to be sensitive to visual disturbance. However, birds are highly intolerant of visual presence and are likely to be scared away by increased human activity, reducing the predation pressure on the mussels. Therefore, visual disturbance may be of indirect benefit to mussel populations. | ||||
Intermediate | High | Low | Moderate | |
Daly & Mathieson (1977) reported that the lower limit of Mytilus edulis populations at Bound Rock, USA, was determined by burial or abrasion by shifting sands. Wave driven logs have been reported to influence Mytilus trossulus (as edulis) populations, causing the removal of patches from extensive beds that subsequently open the beds to further damage by wave action. It is likely that abrasion or impact at the level of the benchmark (a boat anchor being dragged through or landing on the population) would also damage or remove patches of the population. No studies of the effects of trampling on British or Irish populations of Mytilus edulis were found. But the effects of trampling on Mytilus californianus beds in Australia were studied by Brosnan & Cumrine (1994). They exposed mussel beds at two sites to low levels of trampling, 250 steps for 1 day every month over a 1 year period, which compared to 228 steps/hr recorded at another site. They reported that a loose, mono-layer bed was highly susceptible, trampling resulting in patches of bare rock that then expanded, beyond the area trampled, due to wave action. A dense, two layer bed was less susceptible, initially showing less disturbance, although the top layer was lost. However, mussels continued to be lost for a year after trampling had stopped, resulting in patches, and patch size had increased two years after trampling stopped. They suggested that continuous trampling may result in loss of the bed. In a heavily trampled site mussels were not common and were confined to crevices (Brosnan & Cumrine, 1994). Overall, the intolerance of trampling appears to be dependent on the density and depth of the affected mussel bed. Brosnan & Cumrine (1994) observed little recruitment in bare patches in the mussel beds until trampling had ceased, and in some cases no recruitment two years later. Storms and wave action (including wave driven logs) often clear patches of mussels in beds but occur primarily in winter and are localised. Trampling is most likely in spring and summer (Brosnan & Cumrine, 1994). The combined effects of trampling and natural winter disturbances may result in loss of mussel beds in the long term. Mytilus californianus bears radiating ribs, and is a larger than Mytilus edulis with a divergent ecology (Seed, 1992). Nevertheless, the above evidence suggests that mussel beds are potentially intolerant of the effects of trampling, depending on trampling intensity and frequency. Therefore, physical disturbance due to sand abrasion, impact by an anchor or debris, or due to trampling when emersed, is highly likely to result in the loss of a proportion of the population and an intolerance of intermediate has been recorded. Recovery may occur rapidly through good annual recruitment. However, examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported | ||||
Intermediate | High | Low | Moderate | |
Dare (1976) reported that individual mussels swept or displaced from mussel beds rarely survived, since they either became buried in sand or mud, or were scattered and eaten by oystercatchers. Mussels can attach to a wide range of substrata and should a mussel be displaced to a suitable substratum it is likely to be able to attach itself quickly using byssus threads. For example, Young (1985) reported that detached mussels produced 8 byssus threads within the first 24hrs, and between 8-11 byssus threads within 3 days at 13°C (an average of 3.5 threads/ individual/ day), depending on temperature and water agitation. Overall, displacement is likely to result in the loss of some individuals due to vulnerability to predation as well as the danger of smothering, hence an intolerance of intermediate has been recorded. Recovery may occur rapidly through good annual recruitment but examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. |
Intolerance | Recoverability | Sensitivity | Evidence/Confidence | |
Intermediate | High | Low | Moderate | |
The effects of contaminants on Mytilus sp. were extensively reviewed by Widdows & Donkin, (1992) and Livingstone & Pipe (1992). Mussels are suspension feeders and, therefore, process large volumes of water together with suspended particulates and phytoplankton. Mussels absorb contaminants directly from the water, through their diet and via suspended particulate matter (Widdows & Donkin, 1992), the exact pathway being dependant on the nature of the contaminant.
Widdows & Donkin (1992) list tolerances of Mytilus edulis adults and larvae (Tables 8.2, 8.3, & 8.4) but note that lethal responses give a false impression of high tolerance, since the adults can close their valves and isolate themselves from the environment for days. They suggest that sublethal effects (shell growth and 'scope for growth') are more sensitive indicators of the effects of contaminants. Also, adults are ca. 4 times more sensitive than larvae to TBT (Widdows & Donkin, 1992, see larval sensitivity). Overall, the above evidence of contaminant induced mortality suggests that a proportion of the population may be lost and an intolerance of intermediate has been recorded. Recovery may occur rapidly through good annual recruitment but examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. | ||||
Intermediate | High | Low | Low | |
The effects of contaminants on Mytilus sp. were extensively reviewed by Widdows & Donkin, (1992) and Livingstone & Pipe (1992). Widdows & Donkin (1992) list tolerances of Mytilus edulis adults and larvae (Tables 8.2, 8.3, & 8.4) but note that lethal responses give a false impression of high tolerance, since the adults can close their valves and isolate themselves from the environment for days. They suggested that sublethal effects e.g. shell growth and 'scope for growth' (SFG), are more sensitive indicators of the effects of contaminants. Reported effects of heavy metals follow.
Recovery may occur rapidly through good annual recruitment but examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of 'high' has been reported. | ||||
Intermediate | High | Low | Moderate | |
Widdows & Donkin (1992) list tolerances of Mytilus edulis adults and larvae (Tables 8.2, 8.3, & 8.4) but note that lethal responses give a false impression of high tolerance, since the adults can close their valves and isolate themselves from the environment for days. They suggested that sublethal effects e.g. shell growth and 'scope for growth' (SFG), are more sensitive indicators of the effects of contaminants.
Recovery may occur rapidly through good annual recruitment but examination of patches in beds of Mytilus sp. revealed that they may take many years to recover (see additional information below), depending on shore height, competition and environmental conditions. Repeated loss and recruitment results in a patchy distribution of mussels on the shore (Seed & Suchanek, 1992). Therefore, a recoverability of high has been reported. | ||||
No information | Not relevant | No information | Not relevant | |
The periostracum of Mytilus edulis was reported to concentrate uranium (Widdows & Donkin, 1992). Mussels have also been reported to bioaccumulate 106Ru, 95Zr, 95Nb, 137Cs and 90Sr (Cole et al., 1999). While the above data demonstrates that Mytilus edulis can accumulate radionucleides, little information concerning the effects of radionucleides on marine organisms was found. | ||||
Intermediate | High | Low | Low | |
Butler et al. (1990) examined the effects of sewage sludge on adult and larval Mytilus edulis. Exposure of adults to 0.02 - 0.04% industrial/domestic sewage sludge resulted in a reduced respiration rate and a 50% decrease in net energy surplus after 4 weeks. There was no clear relationship between the tissue concentration of heavy metals and physiological stress and it was unclear whether the observed effect was due to increased levels of nutrients or contaminants within the sewage sludge. Mytilus edulis may benefit from moderate nutrient enrichment, especially in the form of organic particulates and dissolved organic material. The resultant increased food availability may increase growth rates, reproductive potential and decrease vulnerability to predators. Long term and/or high levels of organic enrichment may result in deoxygenation (see oxygenation below) and algal blooms, which may have adverse effects indirectly. Algal blooms have reportedly had adverse effects on Mytilus edulis. Blooms of the toxic dinoflagellate Gyrodinium aureolum caused mortality of Mytilus edulis in Norway and sublethal effects on clearance rate and cellular damage in the UK (Holt et al., 1998). A bloom of Phaeocystis poucheri that produced copious amounts of glutinous material prevented Mytilus edulis from feeding, and hence reproductive failure in the Dutch Wadden Sea (Pieters et al., 1980; Holt et al., 1998). Landsberg (1996) also suggested that there was a correlation between the incidence of neoplasia or tumours in bivalves and out-breaks of paralytic shellfish poisoning in which bivalves accumulate toxins from algal blooms, although a direct causal effect required further research. Therefore, a proportion of the population may be lost as a result of nutrient enrichment, and an intolerance of intermediate has been recorded. Recoverability is likely to be high (see additional information below). | ||||
Low | Very high | Very Low | Low | |
Mytilus edulis is likely to encounter hyper-saline conditions in rock pools exposed on hot days, where evaporation can increase the salinity, e.g. Newell (1979) presented data on salinity fluctuations in rock pools, which reached salinities up to 42 psu. High shore rock pools show marked fluctuations in salinity. But Mytilus edulis is considered to be tolerant of a wide range of salinities (see Holt et al., 1998). Therefore, an intolerance of low, at the benchmark level, is recorded. On return to the prior salinity regime, Mytilus edulis will probably recover within a few days or weeks. | ||||
Low | Very high | Very Low | Moderate | |
Mytilus edulis exhibits a defined behaviour to reducing salinity, initially only closing its siphons to maintain the salinity of the water in its mantle cavity, which allows some gaseous exchange and therefore maintains aerobic metabolism for longer. If the salinity continues to fall the valves close tightly (Davenport, 1979; Rankin & Davenport, 1981). In extreme low salinities, e.g. resulting from storm runoff, large numbers of mussels may be killed (Keith Hiscock pers comm.). In the long term (weeks) Mytilus edulis can acclimate to lower salinities (Almada-Villela, 1984; Seed & Suchanek, 1992; Holt et al., 1998). Almada-Villela (1984) reported that the growth rate of individuals exposed to only 13psu reduced to almost zero but had recovered to over 80% of control animals within one month.
Mytilus edulis can also survive considerably reduced salinities, growing as dwarf individuals at 4-5psu in the Baltic. Differences in growth being due to physiological and/or genetic adaptation to salinity.
Mytilus edulis is an osmoconformer and maintains its tissue fluids iso-osmotic (equal ionic strength) with the surrounding medium by mobilization and adjustment of the tissue fluid concentration of free amino acids (e.g. taurine, glycine and alanine) (Bayne, 1976; Newell, 1989). But mobilizing amino acids may result in loss of protein, increased nitrogen excretion and reduced growth. Koehn (1983) and Koehn & Hilbish (1987) reported a genetic basis to adaptation to salinity at the aminopeptidase-1 locus, involved in the production of some free amino acids. In addition, Mytilus edulis thrives in brackish lagoons and estuaries, although, this is probably due to the abundance of food in these environments rather than the salinity (Seed & Suchanek, 1992). Overall, Mytilus edulis can acclimate to a wide range of salinities and a change of salinity at the benchmark level is unlikely to adversely affect this species, and an intolerance of low has been recorded. | ||||
Low | Very high | Very Low | High | |
Mytilus edulis is regarded as euryoxic, tolerant of a wide range of oxygen concentrations including zero (Zwaan de & Mathieu, 1992). Diaz & Rosenberg (1995) suggest it is resistant to severe hypoxia. Adult mytilids exhibited high tolerance of anoxia, e.g. Theede et al. (1969) reported LD50 of 35 days for Mytilus edulis exposed to 0.21mg/l O2 at 10°C, which was reduced to 25 days with the addition of sulphide (50 mg/l Na2S .9H2O). Mytilus edulis is capable of anaerobic metabolism. In aerial exposure (emersion) the mussel closes its valves, resulting in a low rate of oxygen exchange and consumption (Zwaan de & Mathieu, 1992; Widdows et al., 1979). Therefore, the mussel conserves energy and utilizes anaerobic metabolism. Anaerobic metabolism also increases at low temperatures and some of the end products of anaerobic metabolism may be cryoprotectant (see changes in temperature above). Jorgensen (1980) observed, by diving, the effects of hypoxia (0.2 -1 mg/l) on benthic macrofauna in marine areas in Sweden over a 3-4 week period. Mussels were observed to close their shell valves in response to hypoxia and survived for 1-2 weeks before dying (Cole et al., 1999; Jorgensen, 1980). In hypoxic or anoxic conditions Mytilus edulis increases oxygen consumption until oxygen levels fall below 60% saturation, the proportion of anaerobic metabolism increases as the oxygen concentration falls below 90% saturation (Famme et al., 1981; Newell, 1989). In Mytilus galloprovincialis anaerobic metabolism increases once the oxygen concentration falls below 8kPa ( 3.5 mg/l) becoming maximal at and below 4kPa (1.76 mg/l) (Zwaan de & Mathieu, 1992). Anaerobic metabolism allows the mussel to maintain is metabolism close to aerobic levels (Newell, 1989), although it incurs an 'oxygen debt' in the process (Widdows et al., 1979). Although Mytilus edulis is highly tolerant of hypoxia at the benchmark level (2mg/l O2 for 1 week), it incurs a metabolic cost and, hence, reduced growth, therefore, an intolerance of low has been recorded. Tolerance of anoxia increases during larval development (see larval sensitivity). Once oxygen levels return to prior levels, Mytilus edulis will probably recover condition within a few weeks. |
Intolerance | Recoverability | Sensitivity | Evidence/Confidence | |
Intermediate | High | Low | High | |
Landsberg (1996) suggested that there was a correlation between the incidence of neoplasia or tumours in bivalves and out-breaks of paralytic shellfish poisoning in which bivalves accumulate toxins from algal blooms. However, demonstration of a direct causal effect requires further research. Mytilus spp. hosts a wide variety of disease organisms. parasites and commensals from many animal and plant groups including bacteria, blue green algae, green algae, protozoa, boring sponges, boring polychaetes, boring lichen, the intermediary life stages of several trematodes, copepods and decapods (Bower, 1992; Bower & McGladdery, 1996; Gray et al., 1999). For example:
| ||||
No information | Not relevant | No information | Not relevant | |
Mytilus edulis is an effective space occupier and few other species are able to out-compete it for space. However, the South American mytilid Aulocomya ater has been reported recently in the Moray Firth, Scotland in 1994 and again in 1997 (Holt et al., 1998; Eno et al., 2000; McKay, 1994). Aulocomya ater is thought to have a stronger byssal attachment than Mytilus edulis and may replace Mytilus edulis in more exposed areas if it reproduces successfully (Holt et al., 1998). | ||||
Intermediate | High | Low | Moderate | |
Large mussel beds have been routinely fished for hundreds of years, and managed by local Sea Fishery Committees for the past hundred years (Holt et al., 1998). Mussel beds may be exploited by hand collection or dredging. Holt et al., (1998) suggest that when collected by hand at moderate levels using traditional skills the beds will probably retain most of their biodiversity (see importance). However, they also cite incidences of over-exploitation of easily accessible small beds by anglers for bait. Holt et al., (1998) suggest that in particular embayments over-exploitation may reduce subsequent recruitment leading to long term reduction in the population or stock. The relationship between stock and recruitment is poorly understood. Loss of stock may have significant effects on other species, e.g. in the Dutch Wadden Sea in 1990 the mussel stocks fell to unprecedented low levels resulting in death or migration of eiders, and oystercatchers seeking alternative prey such as the cockle Cerastoderma edule, the sand-gaper Mya arenaria and Baltic tellin Macoma balthica. Therefore, extraction or fishing will inevitably result in loss of a proportion of the population so that an intolerance of intermediate has been recorded. Recoverability is probably high (see additional information below). | ||||
Low | Very high | Very Low | Very low | |
Holt et al. (1998) suggested that Mytilus sp. was probably less affected by incidental damage due to fisheries than other organisms, and reported that Mytilus edulis communities replaced Sabellaria spinulosa damaged by shrimp fishing. Therefore, an intolerance of low has been recorded. |
- no data -
National (GB) importance | - | Global red list (IUCN) category | - |
Native | - | ||
Origin | - | Date Arrived | Not relevant |
Ecosystem importance
Larval production represents a significant contribution to the zooplankton, forming an important food source for herring larvae and carnivorous zooplankton (Seed & Suchanek, 1992). Dense beds of bivalve suspension feeders increase turnover of nutrients and organic carbon in estuarine (and presumably coastal) environments by effectively transferring pelagic phytoplanktonic primary production to secondary production (pelagic-benthic coupling) (Dame, 1996).
Nehls & Thiel (1993) suggested that removal or exploitation of persistent mussel beds may reduce or remove food reserves crucial to birds such as eider and oystercatchers. Holt et al. (1998) noted that gross declines in mussel stocks due to over-exploitation or recruitment failure may adversely affect the surrounding ecosystem. For example, in the Dutch Wadden Sea unprecedentedly low mussel numbers in 1990 resulted in death or migration of eider, and oystercatchers seeking alternative prey and increased pressure on cockles, Macoma balthica and Mya arenaria (Holt et al., 1998).
Where farmed shellfish rafts cover 10% or more in areas of poor water exchange (e.g. sea lochs) suspension feeding by mussels can significantly reduce levels of phytoplankton with indirect effects on other filter feeders. Reduced phytoplankton levels in one Scottish loch also adversely affected the farmed mussel production resulting in the closure of the farm. However, most Scottish mussel farms were thought to be too small to have serious effects on plankton levels (McKay & Fowler, 1997; Holt et al., 1998).
Although over-exploitation of local fisheries was identified as a potential problem (Holt et al., 1998) the longevity of the major natural fisheries was probably a result of careful management by Sea Fisheries Committees.
Fisheries
Mussels have been harvested for food and bait since early times. British mussel production is relatively small comprising only 5% of total Europe Community production (Edwards, 1997). Edwards (1997) reported 10,347 tonnes of mussels landed in 1994 with a value of £32 million. Wild mussel fisheries are found in tidal flats of The Wash, Morecambe Bay, Solway and Dornoch Firths in Scotland and river estuaries such as Conwy, North Wales and the Teign and Taw, Devon (Edwards, 1997). Edwards (1997) notes that the commercial development of natural beds is hampered by sporadic and unpredictable recruitment.
Aquaculture
A detailed review of mussel cultivation is provided by Hickman (1992) and the references therein. There has been a move away from exploitation of wild stocks to cultivation in Britain (Edwards, 1997). Examples of cultivation techniques follow.
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Davenport, J., 1979. The isolation response of mussels (Mytilus edulis) exposed to falling sea water concentrations. Journal of the Marine Biological Association of the United Kingdom, 59, 124-132.
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Famme, P., Knudsen, J., & Hansen, E.S., 1981. The effect of oxygen on the aerobic - anaerobic metabolism of the marine bivalve Mytilus edulis. Marine Biology Letters, 2, 345-351.
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This review can be cited as:
Last Updated: 03/06/2008