Polydora ciliata and Corophium volutator in variable salinity infralittoral firm mud or clay

Researched byJacqueline Hill Refereed byThis information is not refereed.
EUNIS CodeA5.321 EUNIS NamePolydora ciliata and Corophium volutator in variable salinity infralittoral firm mud or clay

Summary

UK and Ireland classification

EUNIS 2008A5.321Polydora ciliata and Corophium volutator in variable salinity infralittoral firm mud or clay
EUNIS 2006A5.321Polydora ciliata and Corophium volutator in variable salinity infralittoral firm mud or clay
JNCC 2004SS.SMu.SMuVS.PolCvolPolydora ciliata and Corophium volutator in variable salinity infralittoral firm mud or clay
1997 BiotopeSS.IMU.EstMu.PolVSPolydora ciliata in variable salinity infralittoral firm mud or clay

Description

Variable salinity clay and firm mud characterized by a turf of the polychaete Polydora ciliata. Polydora ciliata also occurs in high densities elsewhere - see MCR.Pol. (May be a specific feature of the Humber Estuary in these conditions). This biotope occurs only in very firm mud and clay and possibly submerged relict saltmarsh with a high detrital content. It is characterized, and can be separated from other biotopes, by a combination of the sediment characteristics and the very high density of Polydora ciliata. (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Recorded distribution in Britain and Ireland

Found in various estuaries along the east coast of Britain, Isle of Wight and the Bristol Channel.

Depth range

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Additional information

None entered

Listed By

Further information sources

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JNCC

Habitat review

Ecology

Ecological and functional relationships

  • In areas of mud, the tubes built by Polydora ciliata can agglomerate and form layers up to an average of 20cm thick, occasionally to 50cm. These mud layers can eliminate the original fauna and flora, or at least can be considered as a threat to the ecological balance achieved by some biotopes (Daro & Polk, 1973).
  • Daro & Polk (1973) state that the formation of layers of Polydora ciliata tend to eliminate original flora and fauna. The species readily overgrows other species with a flat morphology and feeds by scraping its palps outside its tubes, which would inhibit the development of settling larvae of other species.
  • The activities of Polydora plays an important part in the process of temporary sedimentation of muds in some estuaries, harbours or coastal areas (Daro & Polk, 1973).
  • Other species that may be found in the biotope include mud dwelling polychaetes such as the slow moving, burrowing Pholoe synophthalmica and mucus tube building Neoamphitrite figulus. The amphipod Corophium volutator, which builds semi-permanent burrows in mud may also be present in high abundance.

Seasonal and longer term change

  • The early reproductive period of Polydora ciliata often enables the species to be the first in the year to colonize available substrata (Green, 1983). The settling of the first generation in April is followed by the accumulation and active fixing of mud continuously up to a peak during the month of May, when the substrata is covered with the thickest layer of Polydora mud. The following generations do not produce a heavy settlement due to interspecific competition and heavy mortality of the larvae (Daro & Polk, 1973).
  • Later in the year, the surface layer of substantial Polydora muds may not be able to hold the lower layers of the mud mat in place, they crumble away and are then swept away by water currents. The empty tubes of Polydora may saturate the sea close to dense Polydora ciliata aggregations in June. Recolonization of the substratum is made possible, when larva of other species are in the plankton so recolonization by Polydora may not be as successful as earlier in the year.

Habitat structure and complexity

The biotope has very little structural complexity as Polydora tubes aggregate to dominate firm mud or clay. Polydora mats tend to be single species providing little space for other fauna or flora. A Polydora mud is about 20cm thick, but can be up to 50cm thick.

Productivity

  • Productivity in IMU.PolVS is mostly secondary, derived from detritus and organic material. Polydora ciliata generally feeds on detritus but may also suspension feed on particles in the water. Macroalgae are absent from the biotope.
  • The biotope often occurs in nutrient rich areas, for example, close to sewage outfalls. Allochthonous organic material is derived from anthropogenic activity (e.g. sewerage) and natural sources (e.g. plankton, detritus). Autochthonous organic material is formed by benthic microalgae (microphytobenthos e.g. diatoms and euglenoids) and heterotrophic micro-organism production. Organic material is degraded by micro-organisms and the nutrients are recycled. The high surface area of fine particles that covers the Polydora mud provides surface for microflora.

Recruitment processes

The spawning period for Polydora ciliata in northern England is from February until June and three or four generations succeed one another during the spawning period (Gudmundsson, 1985). After a week of development the larvae emerge and are believed to have a pelagic life from two to six weeks before settling (Fish & Fish, 1996). Larvae are substratum specific selecting rocks according to their physical properties or sediment depending on particle size. Larvae of Polydora ciliata have been collected as far as 118km offshore (Murina, 1997). Adults of Polydora ciliata produce a 'mud' resulting from the perforation of soft rock substrata and the larvae of the species settle preferentially on substrata covered with mud (Lagadeuc, 1991).

Time for community to reach maturity

A Polydora biotope is likely to reach maturity very rapidly because Polydora ciliata is a short lived species that reaches maturity within a few months and has three or four spawnings during a breeding season of several months. For example, in colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring. The tubes built by Polydora agglomerate sometimes to form layers of mud up to an average of 20cm thick. However, it may take several years for a Polydora ciliata 'mat' to reach a significant size.

Additional information

None

Preferences & Distribution

Recorded distribution in Britain and IrelandFound in various estuaries along the east coast of Britain, Isle of Wight and the Bristol Channel.

Habitat preferences

Depth Range
Water clarity preferences
Limiting Nutrients Not relevant
Salinity
Physiographic
Biological Zone
Substratum
Tidal
Wave
Other preferences Mud, clay or soft rock substratum

Additional Information

Polydora ciliata is only found in areas of soft rock, such as limestone and chalk, and firm muds and clay where it can make its burrows.

Species composition

Species found especially in this biotope

    Rare or scarce species associated with this biotope

    -

    Additional information

    None

    Sensitivity reviewHow is sensitivity assessed?

    Explanation

    Polydora ciliata is used to assess the sensitivity of the biotope. There has been some confusion in the identification of Polydora species and it has been suggested that some other species such as Polydora ligni, Polydora websteri, Polydora cirrosa and Polydora nuchalis may only be varieties of Polydora ciliata (Mustaquim, 1986). Since no or very few other species are present in the biotope the sensitivity of Polydora ciliata is representative of the sensitivity of the whole biotope.

    Species indicative of sensitivity

    Community ImportanceSpecies nameCommon Name
    Key functionalPolydora ciliataA bristle worm

    Physical Pressures

     IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
    High High Moderate Major decline High
    Removal of the substratum, perhaps by dredging, would result in the loss of Polydora ciliata tubes and hence the loss of the animals so intolerance is assessed as high. However, if some individuals remain, rapid recolonization of the remaining substratum is possible because the species is capable of tube building throughout its life. For example when Daro & Polk (1973) removed Polydora ciliata individuals of all ages on several occasions, all built new tubes. Recovery is likely to be high because the larvae of Polydora ciliata are planktonic and capable of dispersal over long distances and the reproductive period is of several months duration. In colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring.
    Tolerant Not relevant Not relevant No change Moderate
    Polydora ciliata is probably relatively tolerant smothering by 5 cm of sediment because the species inhabits a range of habitats including muddy sediment, larvae settle preferentially on substrates covered with mud (Lagadeuc, 1991) and worms can rebuild tubes close to the surface. The species also plays an important part in the process of temporary sedimentation of muds in some estuaries, harbours or coastal areas (Daro & Polk, 1973). Adults of Polydora ciliata produce a 'mud' resulting from the burrowing and tube building activities in mud, clay and soft rock substrata (Lagadeuc, 1991). A Polydora mud can be up to 50cm thick, but the animals themselves occupy only the first few centimetres. They either elongate their tubes, or leave them to rebuild tubes close to the surface.
    Tolerant Not relevant Not relevant No change High
    Polydora ciliata is able to inhabit a wide range of habitats from muddy sediments to soft rock. For example, the species is found in turbid waters with high levels of suspended sediment which it actively fixes in the process of tube making. Daro & Polk (1973) report that the success of Polydora is directly related to the quantities of muds of any origin carried along by rivers or coastal current. In the Firth of Forth Polydora ciliata formed extensive mats in areas that had an average of 68mg/l suspended solids and a maximum of approximately 680mg/l indicating the species is able to tolerate a huge range of quantities of suspended solids (Read et al., 1982; Read et al., 1983). Occasionally, in certain places siltation is speeded up when Polydora ciliata is present because the species actually produces a 'mud' as it perforates soft rock and chalk habitats and larvae settle preferentially on substrates covered with mud (Lagadeuc, 1991). Therefore, it seems likely that the biotope will be relatively tolerant of increases in suspended sediment and siltation.
    Low High Not relevant No change Low
    Polydora ciliata is able to inhabit a wide range of habitats from muddy sediments to soft rock. Occasionally, in certain places siltation is speeded up when Polydora ciliata is present. Suspended sediment and siltation of those particles is important for tube building in Polydora ciliata so a decrease may reduce tube building or the thickness of the mud surrounding the 'colonies'. Daro & Polk (1973) report that the success of Polydora is directly related to the quantities of muds of any origin carried along by rivers or coastal currents. However, at the level of the benchmark the effects are not likely to be significant and an intolerance rank of low is recorded.
    Not relevant Not relevant Not relevant Not relevant High
    The biotope only occurs in the infralittoral zone and so is not subject to desiccation. Therefore, the factor is not relevant.
    Not relevant Not relevant Not relevant Not relevant High
    The biotope only occurs in the infralittoral zone and so will not be subject to an increase in emergence. Therefore, the factor is not relevant.
    Not sensitive* High
    The biotope only occurs in the infralittoral zone and so is not subject to a decrease in emergence. Therefore, the factor is not relevant.
    Intermediate Very high Low No change Moderate
    Polydora ciliata was present and colonized test panels in Helgoland in three areas, two exposed to strong tidal currents and one site sheltered from currents (Harms & Anger, 1983) so the species appears to tolerate a wide range of water flow regimes. However, in very strong tidal currents little sediment deposition will take place resulting in coarse sediments retaining little organic matter and may become unsuitable for the deposit feeding and tube building activities of Polydora ciliata. Nevertheless, where suspended sediment levels are high, deposition of fine sediment may occur even in strong flows providing suitable conditions for the species. Very strong water flows may sweep away Polydora colonies, often in a thick layer of mud on a hard substratum. If the species tube is embedded in a burrow excavated in limestone rock, shells or calcareous algae the animals may be protected from being washed away in increased flow. However, a change in water flow of 2 categories (see benchmark) for a period of a year is likely to interfere with feeding and tube building by removing sediments and may wash some individuals away. The viability of the biotope is likely to be reduced and so intolerance is set at intermediate. Recovery is very high because the larvae of Polydora ciliata are planktonic and capable of dispersal over long distances and the reproductive period is of several months duration. In colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring.
    Low Very high Moderate No change Moderate
    Polydora ciliata was present and colonized test panels in Helgoland in three areas, two exposed to strong tidal currents and one site sheltered from currents (Harms & Anger, 1983) so the species appears to tolerate a wide range of water flow regimes. A decrease in water flow rate may reduce the suspended particulate material carried in the water column important for Polydora ciliata tube building and feeding. Thus effects will be sub-lethal so intolerance is assessed as low. On return to normal conditions recovery will be very high because Polydora ciliata is able to rapidly recolonize suitable substrata.
    Low Very high Very Low No change Moderate
    Murina (1997) categorised Polydora ciliata as a eurythermal species because of its ability to spawn in temperatures ranging from 10.6-19.9°C. This is consistent with a wide distribution in north-west Europe which extends into the warmer waters of Portugal and Italy (Pardal et al., 1993; Sordino et al., 1989). In the western Baltic Sea Gulliksen (1977) recorded high abundances of Polydora ciliata in temperatures of 7.5 to 11.5°C and in Whitstable in Kent sea temperatures varied between 0.5 and 17°C (Dorsett, 1961). Although there was no information found on the maximum temperature tolerated by Polydora ciliata it does seem likely that the species is able to tolerate a long term increase in temperature of 2°C and may tolerate a short term increase of 5°C. However, growth rates may increase if temperature rises. For example, at Whitstable in Kent, Dorsett (1961) found that a rapid increase in growth coincided with the rising temperature of the sea water during March. However, since effects are likely to be sub-lethal, the intolerance of the biotope is assessed as low. Recovery of the species will be very high because growth and fecundity will return to normal when conditions become more favourable.
    Low Very high Moderate No change High
    Murina (1997) categorised Polydora ciliata as a eurythermal species because of its ability to spawn in temperatures ranging from 10.6-19.9°C. This is consistent with a wide distribution in north-west Europe. In the western Baltic Sea, Gulliksen (1977) recorded high abundances of Polydora ciliata in temperatures of 7.5 to 11.5°C and in Whitstable in Kent abundance was high when winter water temperatures dropped to 0.5°C (Dorsett, 1961). Rapid changes in hydrographical conditions occurred when temperatures dropped from 11.5°C to 7.5°C in the course of 15 hours (Gulliksen, 1977) and so it appears the species is tolerant of short term changes in temperature. During the extremely cold winter of 1962/63 Polydora ciliata was apparently unaffected (Crisp (ed.), 1964). However, it is likely that growth and fecundity may be affected. The intolerance of the biotope is therefore assessed as low because Polydora ciliata appears to be tolerant of both long and short term decreases in temperature. The high fecundity, short generation times and large dispersal distances achieved by Polydora ciliata means the species, and hence the biotope, will probably recover very rapidly on return to normal conditions.
    Low Very high Very Low No change High
    An increase in turbidity, reducing light availability, may reduce primary production by phytoplankton in the water column. However, productivity in the MCR.Pol biotope is secondary because Polydora ciliata deposit feeds on detritus or suspended particles in the water column. Therefore, the biotope is not likely to be significantly affected by changes in turbidity and so intolerance is assessed as low. In estuaries and surf zones on the lower shore, turbidity can be measured in g/l so the benchmark level is low in comparison. Nevertheless, primary production by pelagic phytoplankton and microphytobenthos do contribute to benthic communities and so long term increases in turbidity may reduce the overall organic input to the detritus. Reduced food supply may affect growth rates and fecundity of some species in the biotope. However, at the level of the benchmark effects are not likely to be significant and a rank of low intolerance is reported. On return to normal turbidity levels recovery will be very high as food availability returns to normal.
    Low Very high Moderate No change High
    A decrease in turbidity, increasing light availability may increase primary production by phytoplankton in the water column. However, productivity in the MCR.Pol biotope is secondary because Polydora ciliata deposit feeds on detritus or suspended particles in the water column. Therefore, the biotope is not likely to be significantly affected by changes in turbidity and so intolerance is assessed as low. Nevertheless, primary production by pelagic phytoplankton and microphytobenthos do contribute to benthic communities and long term decreases in turbidity may increase the overall organic input to the detritus. Increased food supply may increase growth rates and fecundity of some species in the biotope.
    High Very high Low No change Moderate
    The biotope is found in a range of wave exposures from very sheltered to moderately exposed. Feeding may be impaired in strong wave action and changes in wave exposure may also influence the supply of particulate matter. Polydora tubes normally form into 'mats' which are likely to be washed away if exposure were to increase by two exposure scales for a year and so intolerance is reported to be high.
    Low Very high Moderate No change Moderate
    The biotope is found in moderately wave exposed to very wave sheltered sites. A decrease in wave exposure may influence the supply of particulate matter for suspension feeding because wave action may have an important role in re-suspending the sediment that is required by the species to build its tubes. Food supplies may also be reduced affecting growth and fecundity of the species. Abundance of the species may decline if wave exposure decreases. However, since the benchmark level is likely to result in sub-lethal changes only the intolerance of the biotope is regarded to be low.
    Tolerant Not relevant Not relevant No change Moderate
    Polydora ciliata may respond to vibrations from predators or bait diggers by retracting their palps into their tubes. However, the species is unlikely to sensitive to noise at the benchmark levels and so the biotope is assessed as not sensitive.
    Low Immediate Not sensitive No change Moderate
    Polydora ciliata exhibits shadow responses withdrawing its palps into its burrow, believed to be a defence against predation. However, since the withdrawal of the palps interrupts feeding and possibly respiration the species also shows habituation of the response (Kinne, 1970). The species is, therefore, likely to have very low intolerance to visual disturbance and the biotope will be little affected by the presence of boats, humans or other visual factors not normally present in the marine environment.
    Intermediate High Low No change Moderate
    A passing scallop dredge (or other mobile fishing gear) is likely to remove or break up the tubicolous mat of Polydora ciliata. Individuals are likely to be crushed and killed. However, some individuals are likely to survive, as individuals are small and can withdraw into burrows. Therefore, intolerance has been assessed as intermediate. Recovery is good because Polydora ciliata has planktonic larvae that are capable of dispersal over long distances and the reproductive period is of several months duration. In colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring.
    Low High Low No change High
    Polydora ciliata is capable of tube building throughout its life and so is able to re-establish attachment on displacement. For example, in removal experiments, Polydora ciliata individuals of all ages that were removed from their tubes on many occasions built new tubes (Daro & Polk, 1973). Recovery is likely to be high because Polydora ciliata has planktonic larvae that are capable of dispersal over long distances and the reproductive period is of several months duration. In colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring.

    Chemical Pressures

     IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
    Low High Low No change Not relevant
    The biotope occurs in polluted sites and so has a low intolerance to the factor. For example, Polydora ciliata was abundant at polluted sites close to acidified, halogenated effluent discharge from a bromine-extraction plant in Amlwch, Anglesey (Hoare & Hiscock, 1974). Spionid polychaetes were found by McLusky (1982) to be relatively tolerant of distilling and petrochemical industrial waste in Scotland.
    Heavy metal contamination
    Intermediate Very high Low No change Moderate
    Experimental studies with various species suggests that polychaete worms are quite tolerant to heavy metals (Bryan, 1984). Polydora ciliata occurs in an area of the southern North Sea polluted by heavy metals but was absent from sediments with very high heavy metal levels (Diaz-Castaneda et al., 1989). Intolerance has thus been assessed to be intermediate. Recovery is likely to be very high - see additional information below.
    Hydrocarbon contamination
    Intermediate Very high Low No change Moderate
    In analysis of kelp holdfast fauna following the Sea Empress oil spill in Milford Haven the fauna present, including Polydora ciliata, showed a strong negative correlation between numbers of species and distance from the spill (SEEEC, 1998). After the extensive oil spill in West Falmouth, Massachusetts, Grassle & Grassle (1974) followed the settlement of polychaetes in this environmental disturbed area. Species with the most opportunistic life histories, including Polydora ligni, were able to settle in the area. Intolerance has therefore, been assessed as intermediate. This species has some brood protection which enables larvae to settle almost immediately in the nearby area (Reish, 1979) and so recovery is expected to be very high - see below for additional information.
    Radionuclide contamination
    No information No information No information Insufficient
    information
    Not relevant
    Insufficient
    information.
    Changes in nutrient levels
    Tolerant* Very high Not sensitive* No change High
    Polydora ciliata is often found in environments subject to high levels of nutrients. For example, the species was abundant in areas of the Firth of Forth exposed to high levels of sewage pollution (Smyth, 1968) and in nutrient rich sediments in the Mondego estuary, Portugal (Pardal et al., 1993) and the coastal lagoon Lago Fusaro in Naples (Sordino et al., 1989). Extensive growths of Polydora ciliata in mat formations were recorded at West Ganton, in the Firth of Forth, prior to the introduction of a sewage treatment scheme (Read et al., 1983). The abundance of the species was probably associated with their ability to use the increased availability of organic matter as a food source and silt for tube building. As water quality improved following introduction of the scheme these 'pollution tolerant' species disappeared providing space for colonization by other fauna (Read et al., 1983). Polydora ciliata can also occur in organically poor areas (Pearson & Rosenberg, 1978). The species and hence the biotope is likely to benefit from increased nutrients and so a rank of 'not sensitive*' is recorded. This response is likely to be very rapid. For example, in colonization experiments in an organically polluted fjord receiving effluent discharge from Oslo, Polydora ciliata had settled in large numbers within the first month (Green, 1983, Pardal et al., 1993) and in colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring.
    High High Moderate No change Low
    Polydora ciliata is a euryhaline species inhabiting fully marine and estuarine habitats. However, there are no records of the species or the biotope occurring in hypersaline waters and an increase for a period of a year is likely to result in the death of many individuals and so intolerance is reported to be high.
    Low High Low No change Low
    Polydora ciliata is a euryhaline species inhabiting fully marine and estuarine habitats. In an area of the western Baltic Sea, where bottom salinity was between 11.1 and 15.0psu Polydora ciliata was the second most abundant species with over 1000 individuals per m2 (Gulliksen, 1977). Intolerance to a decrease in salinity is therefore, expected to be low.
    Tolerant* High Not sensitive* No change High
    Polydora ciliata has low intolerance to reduced oxygenation because the species is repeatedly found at localities with oxygen deficiency (Pearson & Rosenberg, 1978). For example, in polluted waters in Los Angeles and Long Beach harbours Polydora ciliata was present in the oxygen range 0.0-3.9 mg/l and the species was abundant in hypoxic fjord habitats (Rosenberg, 1977). Only in very hypoxic and anoxic conditions can Polydora ciliata not survive. Therefore, at the benchmark level of deoxygenation the biotope may establish or expand as a result of reduced oxygen adversely affecting other species and so a rank of 'not sensitive* is indicated.

    Biological Pressures

     IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
    No information No information No information Insufficient
    information
    Not relevant
    No information on diseases affecting Polydora ciliata or the biotope was found.
    Tolerant Not relevant Not relevant Not relevant Moderate
    No known non-native species compete with Polydora ciliata and so the biotope is assessed as not sensitive.
    Not relevant Not relevant Not relevant Not relevant Not relevant
    The biotope consists of Polydora ciliata to the exclusion of almost all other species. Extraction of Polydora ciliata is unlikely and we have no evidence for the indirect effects of extraction of other species on this biotope. However, if dredging occurred, populations may be removed in some habitats (see Physical Disturbance above). Overall this factor is not considered to be relevant.
    Not relevant Not relevant Not relevant Not relevant Moderate

    Additional information

    Recoverability
    Effects on the biotope, even significant effects, will not change the species diversity because the biotope is defined as a single species community.

    Importance review

    Policy/Legislation

    Habitats Directive Annex 1Estuaries

    Exploitation

    The biotope is unlikely to be exploited because Polydora ciliata is of no commercial interest.

    Additional information

    -

    Bibliography

    1. Daro, M.H. & Polk, P., 1973. The autecology of Polydora ciliata along the Belgian coast. Netherlands Journal of Sea Research, 6, 130-140.
    2. Diaz-Castaneda, V., Richard, A. & Frontier, S., 1989. Preliminary results on colonization, recovery and succession in a polluted areas of the southern North Sea (Dunkerque's Harbour, France). Scientia Marina, 53, 705-716.
    3. Dorsett, D.A., 1961. The reproduction and maintenance of Polydora ciliata (Johnst.) at Whitstable. Journal of the Marine Biological Association of the United Kingdom, 41, 383-396.
    4. Fish, J.D. & Fish, S., 1974. The breeding cycle and growth of Hydrobia ulvae in the Dovey estuary. Journal of the Marine Biological Association of the United Kingdom, 54, 685-697.
    5. Grassle, J.F. & Grassle, J.P., 1974. Opportunistic life histories and genetic systems in marine benthic polychaetes. Journal of Marine Research, 32, 253-284.
    6. Green, J., 1961. A biology of Crustacea. London: H.F. & G. Witherby Ltd. 180pp.
    7. Green, N.W., 1983. Key colonisation strategies in a pollution-perturbed environment. In Fluctuations and Succession in Marine Ecosystems: Proceedings of the 17th European Symposium on Marine Biology, Brest, France, 27 September - 1st October 1982. Oceanologica Acta, 93-97.
    8. Gudmundsson, H., 1985. Life history patterns of polychaete species of the family spionidae. Journal of the Marine Biological Association of the United Kingdom, 65, 93-111.
    9. Gulliksen, B., 1977. Studies from the U.W.L. "Helgoland" on the macrobenthic fauna of rocks and boulders in Lübeck Bay (western Baltic Sea). Helgoländer wissenschaftliche Meeresunters, 30, 519-526.
    10. Harms, J. & Anger, K., 1983. Seasonal, annual, and spatial variation in the development of hard bottom communities. Helgoländer Meeresuntersuchungen, 36, 137-150.
    11. Hoare, R. & Hiscock, K., 1974. An ecological survey of the rocky coast adjacent to the effluent of a bromine extraction plant. Estuarine and Coastal Marine Science, 2 (4), 329-348.

    12. Kinne, O. (ed.), 1970. Marine Ecology: A Comprehensive Treatise on Life in Oceans and Coastal Waters. Vol. 1 Environmental Factors Part 1. Chichester: John Wiley & Sons
    13. Lagadeuc, Y., 1991. Mud substrate produced by Polydora ciliata (Johnston, 1828) (Polychaeta, Annelida) - origin and influence on fixation of larvae. Cahiers de Biologie Marine, 32, 439-450.
    14. McLusky, D.S., 1982. The impact of petrochemical effluent on the fauna of an intertidal estuarine mudflat. Estuarine, Coastal and Shelf Science, 14, 489-499.
    15. Murina, V., 1997. Pelagic larvae of Black Sea Polychaeta. Bulletin of Marine Science, 60, 427-432.
    16. Mustaquim, J., 1986. Morphological variation in Polydora ciliata complex (Polychaeta, Annelida). Zoological Journal of the Linnean Society, 86, 75-88.
    17. Pardal, M.A., Marques, J.-C. & Bellan, G., 1993. Spatial distribution and seasonal variation of subtidal polychaete populations in the Mondego estuary (western Portugal). Cahiers de Biologie Marine, 34, 497-512.
    18. Pearson, T.H. & Rosenberg, R., 1978. Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. Oceanography and Marine Biology: an Annual Review, 16, 229-311.
    19. Read, P.A., Anderson, K.J., Matthews, J.E., Watson, P.G., Halliday, M.C. & Shiells, G.M., 1982. Water quality in the Firth of Forth. Marine Pollution Bulletin, 13, 421-425.
    20. Read, P.A., Anderson, K.J., Matthews, J.E., Watson, P.G., Halliday, M.C. & Shiells, G.M., 1983. Effects of pollution on the benthos of the Firth of Forth. Marine Pollution Bulletin, 14, 12-16.
    21. Reish, D.J., 1979. Bristle Worms (Annelida: Polychaeta) In Pollution Ecology of Estuarine Invertebrates, (eds. Hart, C.W. & Fuller, S.L.H.), 78-118. Academic Press Inc, New York.
    22. SEEEC (Sea Empress Environmental Evaluation Committee), 1998. The environmental impact of the Sea Empress oil spill. Final Report of the Sea Empress Environmental Evaluation Committee, 135 pp., London: HMSO.
    23. Smyth, J.C., 1968. The fauna of a polluted site in the Firth of Forth. Helgolander Wissenschaftliche Meeresuntersuchungen, 17, 216-233.
    24. Sordino, P., Gambi, M.C. & Carrada, G.C., 1989. Spatio-temporal distribution of polychaetes in an Italian coastal lagoon (Lago Fusaro, Naples). Cahiers de Biologie Marine, 30, 375-391.

    Citation

    This review can be cited as:

    Hill, J.M. 2002. Polydora ciliata and Corophium volutator in variable salinity infralittoral firm mud or clay. 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/habitat/detail/193

    Last Updated: 25/11/2002