A bristleworm (Polydora ciliata)
Distribution data supplied by the Ocean Biodiversity Information System (OBIS). To interrogate UK data visit the NBN Atlas.Map Help
Researched by | Jacqueline Hill | Refereed by | This information is not refereed |
Authority | (Johnston, 1838) | ||
Other common names | - | Synonyms | - |
Summary
Description
A sedentary, burrowing polychaete worm up to 3 cm long and 0.7-1 mm wide. The body has up to 180 segments but is not divided into distinct regions. Polydora ciliata has two very long, slender ciliated palps which protrude, waving vigorously and usually roll up spirally when the animal is disturbed. The tip of the posterior region is saucer-shaped. Polydora ciliata is yellowish-brown in colour.
Recorded distribution in Britain and Ireland
Polydora ciliata is widely distributed around Britain and Ireland.Global distribution
Widely distributed in north-west Europe.Habitat
Polydora ciliata usually burrows into substrata containing calcium carbonate such as limestone and chalk and into clay as well as the shells of oysters, mussels and periwinkles and crusts of calcareous algae ('lithothamnia'). The species is also found in muddy sediments, wood and laminarian holdfasts.Depth range
-Identifying features
- Prostomium blunt in front, extending rearwards in a low crest.
- 5th chaetiger (segment) is enlarged and different from all the others, lacking gills and parapodial lobes but bearing six or seven extra-large chaetal spines, with a lateral tooth, dorsally.
- Gills present on dorsal surface from chaetiger 7 to all but the ten rearmost ones.
- Short, stout chaetae ventrally from seventh or eighth chaetiger, tipped with tow small hooded teeth; this type of chaeta absent dorsally.
- Longitudinal sensory grooves mid-dorsally.
- Anus surrounded by a membranous funnel.
- Number of eyes varies from zero to four (Mustaquim, 1986).
Additional information
There has been some confusion in the identification of Polydora ciliata because the characteristics used for separation of the species, such as the number of modified chaetae on the fifth segment, are not stable even in individuals from the same locality. It has been suggested that some other species of Polydora such as P. ligni, P. websteri, P. cirrosa and P. nuchalis may only be varieties of Polydora ciliata (Mustaquim, 1986).
Listed by
- none -
Biology review
Taxonomy
Level | Scientific name | Common name |
---|---|---|
Phylum | Annelida | Segmented worms e.g. ragworms, tubeworms, fanworms and spoon worms |
Class | Polychaeta | Bristleworms, e.g. ragworms, scaleworms, paddleworms, fanworms, tubeworms and spoon worms |
Order | Spionida | |
Family | Spionidae | |
Genus | Polydora | |
Authority | (Johnston, 1838) | |
Recent Synonyms |
Biology
Parameter | Data | ||
---|---|---|---|
Typical abundance | High density | ||
Male size range | |||
Male size at maturity | |||
Female size range | Small(1-2cm) | ||
Female size at maturity | |||
Growth form | Vermiform segmented | ||
Growth rate | |||
Body flexibility | High (greater than 45 degrees) | ||
Mobility | |||
Characteristic feeding method | Active suspension feeder, Surface deposit feeder | ||
Diet/food source | |||
Typically feeds on | Detritus | ||
Sociability | |||
Environmental position | Epibenthic | ||
Dependency | Independent. See additional information | ||
Supports | None | ||
Is the species harmful? | No information |
Biology information
Mode of life. Polydora ciliata burrows into the shells of oysters, mussels and periwinkles as well as into limestone rock and stones and lithothamnia or other encrusting coralline algae. The species makes a U-shaped tube from small particles (usually of mud) but may be whitish and calcareous if excavating in lithothamnia or other encrusting coralline algae (Hayward & Ryland, 1995). Much of this tube may be embedded in a burrow excavated in limestone rock, shells and calcareous algae, and the two ends extend a few millimetres above the surface of the substratum. It has been suggested that burrowing is achieved by the mechanical action of the chaetae, especially those of the 5th segment, but this is open to some doubt as chemical action may also be involved (Fish & Fish, 1996).
Feeding method. The species generally feeds on detritus that is removed from the sediment by the two long palps. It also feeds on suspended particles in the water, and on occasions has been observed to eat dead barnacles and other dead invertebrates.
Habitat preferences
Parameter | Data |
---|---|
Physiographic preferences | Open coast, Offshore seabed, Strait or Sound, Estuary, Isolated saline water (Lagoon), Enclosed coast or Embayment |
Biological zone preferences | Lower circalittoral, Lower eulittoral, Lower infralittoral, Mid eulittoral, Sublittoral fringe, Upper circalittoral, Upper infralittoral |
Substratum / habitat preferences | Macroalgae, Artificial (man-made), Bedrock, Mud, Other species |
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, Extremely sheltered, Moderately exposed, Sheltered, Very sheltered |
Salinity preferences | Full (30-40 psu), Low (<18 psu), Variable (18-40 psu) |
Depth range | |
Other preferences | No text entered |
Migration Pattern | Non-migratory or resident |
Habitat Information
The species makes a U-shaped tube from small particles (usually of mud) but may be whitish and calcareous if excavating in lithothamnia or other encrusting coralline algae (Hayward & Ryland, 1995). Much of this tube may be embedded in a burrow excavated in limestone rock, shells and calcareous algae, and the two ends extend a few millimetres above the surface of the substratum. It has been suggested that burrowing is achieved by mechanical action of the chaete, especially those of the 5th segment, but this is open to some doubt as chemical action may also be involved (Fish & Fish, 1996).Life history
Adult characteristics
Parameter | Data |
---|---|
Reproductive type | Gonochoristic (dioecious) |
Reproductive frequency | Annual protracted |
Fecundity (number of eggs) | 1,000-10,000 |
Generation time | <1 year |
Age at maturity | 2-3 months |
Season | February - June |
Life span | <1 year |
Larval characteristics
Parameter | Data |
---|---|
Larval/propagule type | - |
Larval/juvenile development | Planktotrophic |
Duration of larval stage | See additional information |
Larval dispersal potential | Greater than 10 km |
Larval settlement period | Insufficient information |
Life history information
- Sperm are drawn into the burrow of the female in the respiratory current and the eggs are laid in a string of capsules. A single female produces many capsules, each containing up to about 60 eggs, the individual capsules being attached by two threads to the wall of the burrow. Capsules are brooded for about a week before the larvae are released into the water column.
- Spawning period varies, from February until June in northern England (Gudmundsson, 1985) and in the Black Sea spawning lasted from April - September (Murina, 1997). In Belgium (Daro & Polk, 1973) and northern England (Gudmundsson, 1985) three or even four generations succeeded one another during the spawning period. The number of offspring produced per female varied from 200 to 2200.
- After a week, the larvae emerge and are believed to have a pelagic life from two to six weeks before settling (Fish & Fish, 1996). Settlement and metamorphosis takes place when the larvae has 17-18 setigers.
- Larvae are substratum specific selecting rocks according to their physical properties or sediment depending on substrate particle size.
- Larvae of Polydora ciliata have been collected as far as 118km offshore (Murina, 1997) and along the Belgian coast were found in the plankton all year round with a peak in the summer (Daro & Polk 1973).
Sensitivity review
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.
Physical pressures
Use / to open/close text displayed
Intolerance | Recoverability | Sensitivity | Evidence / Confidence | |
Substratum loss [Show more]Substratum lossBenchmark. All of the substratum occupied by the species or biotope under consideration is removed. A single event is assumed for sensitivity assessment. Once the activity or event has stopped (or between regular events) suitable substratum remains or is deposited. Species or community recovery assumes that the substratum within the habitat preferences of the original species or community is present. Further details EvidenceRemoval 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 is possible because the species is capable of tube building throughout its life. Polydora ciliata of all ages that were removed from their tubes on many occasions, all built new tubes (Daro & Polk, 1973). 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. | High | High | Moderate | Moderate |
Smothering [Show more]SmotheringBenchmark. All of the population of a species or an area of a biotope is smothered by sediment to a depth of 5 cm above the substratum for one month. Impermeable materials, such as concrete, oil, or tar, are likely to have a greater effect. Further details. EvidencePolydora ciliata is likely to tolerate smothering by 5 cm of sediment because the species inhabits a range of habitats including muddy sediment and larvae settle preferentially on substrates covered with mud (Lagadeuc, 1991). 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). 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 have left them to rebuild close to the surface. | Tolerant | Not relevant | Not sensitive | Moderate |
Increase in suspended sediment [Show more]Increase in suspended sedimentBenchmark. An arbitrary short-term, acute change in background suspended sediment concentration e.g., a change of 100 mg/l for one month. The resultant light attenuation effects are addressed under turbidity, and the effects of rapid settling out of suspended sediment are addressed under smothering. Further details EvidencePolydora ciliata is tolerant to siltation because it normally inhabits waters with high levels of suspended sediment which it actively fixes in the process of tube making when in muddy habitats. Occasionally, in certain places siltation is speeded up when Polydora ciliata is present. | Tolerant | Not relevant | Not sensitive | High |
Decrease in suspended sediment [Show more]Decrease in suspended sedimentBenchmark. An arbitrary short-term, acute change in background suspended sediment concentration e.g., a change of 100 mg/l for one month. The resultant light attenuation effects are addressed under turbidity, and the effects of rapid settling out of suspended sediment are addressed under smothering. Further details Evidence | No information | |||
Desiccation [Show more]Desiccation
EvidencePolydora ciliata colonizes a wide range of littoral and sub-littoral habitats from rocks on the midshore to the subtidal and are therefore tolerant to a level of desiccation. For example, at Cullercoats in north east England, animals were present at the mid-shore level, where the worms are subjected to about equal time of exposure and submergence (Gudmundsson, 1985). Although soft bodies are likely to be intolerant of desiccation Polydora ciliata can retreat into its burrow to ameliorate the effects. Therefore, only those individuals at the upper limit of the population range are likely to be killed by an increase in desiccation and so intolerance has been assessed as intermediate. | Intermediate | High | Low | Moderate |
Increase in emergence regime [Show more]Increase in emergence regimeBenchmark. A one hour change in the time covered or not covered by the sea for a period of one year. Further details EvidencePolydora ciliata colonizes a wide range of littoral and sub-littoral habitats from rocks on the midshore to the subtidal and are therefore tolerant to a level of emergence. For example, at Cullercoats in north east England, animals were present at the mid-shore level, where they are subjected to about equal time of exposure and submergence (Gudmundsson, 1985). An increase in emergence may cause the death of some individuals at the upper limit of the species range because of increased desiccation. Sub-tidal populations are unlikely to experience emergence. | Intermediate | High | Low | Moderate |
Decrease in emergence regime [Show more]Decrease in emergence regimeBenchmark. A one hour change in the time covered or not covered by the sea for a period of one year. Further details Evidence | No information | |||
Increase in water flow rate [Show more]Increase in water flow rateA change of two categories in water flow rate (view glossary) for 1 year, for example, from moderately strong (1-3 knots) to very weak (negligible). Further details EvidencePolydora 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). In very strong tidal currents little sediment deposition will take place resulting in coarse sediments retaining little organic matter and therefore, not suitable for the deposit feeding Polydora ciliata. However, where suspended sediment levels are high, deposition of fine sediment may occur even in strong flows providing suitable conditions for the species. Animals living in burrows in rock are not likely to be washed away but strong water flow rate may interfere with feeding and tube building by removing sediments. Recovery is good because animals can re-build tubes and 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. | Intermediate | High | Low | Moderate |
Decrease in water flow rate [Show more]Decrease in water flow rateA change of two categories in water flow rate (view glossary) for 1 year, for example, from moderately strong (1-3 knots) to very weak (negligible). Further details Evidence | No information | |||
Increase in temperature [Show more]Increase in temperature
For intertidal species or communities, the range of temperatures includes the air temperature regime for that species or community. Further details EvidenceMurina (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 global distribution. In the western Baltic Sea Gulliksen (1977) recorded high abundances of Polydora ciliata in temperatures of 7.5 to 11.5°C. 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 when temperatures dropped below freezing point for several weeks, Polydora ciliata was apparently unaffected (Crisp (ed.), 1964). Recovery of the species is good because the larvae of Polydora ciliata are planktonic and capable of dispersal over long distances and the reproductive period is of several months duration. | Low | High | Low | Moderate |
Decrease in temperature [Show more]Decrease in temperature
For intertidal species or communities, the range of temperatures includes the air temperature regime for that species or community. Further details Evidence | No information | |||
Increase in turbidity [Show more]Increase in turbidity
EvidenceThe species is probably tolerant of changes in turbidity because it is able to colonize a range of habitats including muddy sediments and soft rock substratum that vary in turbidity. | Tolerant | Not relevant | Not sensitive | Moderate |
Decrease in turbidity [Show more]Decrease in turbidity
Evidence | No information | |||
Increase in wave exposure [Show more]Increase in wave exposureA change of two ranks on the wave exposure scale (view glossary) e.g., from Exposed to Extremely exposed for a period of one year. Further details EvidenceIn the intertidal, Polydora ciliata generally inhabits a burrow within rocks and so is unlikely to be damaged or removed by exposure to wave action and so intolerance is assessed as low. Changes in wave exposure may influence the supply of particulate matter for suspension feeding. | Low | High | Low | Low |
Decrease in wave exposure [Show more]Decrease in wave exposureA change of two ranks on the wave exposure scale (view glossary) e.g., from Exposed to Extremely exposed for a period of one year. Further details Evidence | No information | |||
Noise [Show more]Noise
EvidencePolydora ciliata may respond to vibrations from predators or bait diggers by retracting their palps into their tubes. However, the species is unlikely to be sensitive to noise. | Tolerant | Not relevant | Not sensitive | Moderate |
Visual presence [Show more]Visual presenceBenchmark. The continuous presence for one month of moving objects not naturally found in the marine environment (e.g., boats, machinery, and humans) within the visual envelope of the species or community under consideration. Further details EvidencePolydora 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 by boats, humans or other factors not normally present in the marine environment. | Low | High | Low | Moderate |
Abrasion & physical disturbance [Show more]Abrasion & physical disturbanceBenchmark. Force equivalent to a standard scallop dredge landing on or being dragged across the organism. A single event is assumed for assessment. This factor includes mechanical interference, crushing, physical blows against, or rubbing and erosion of the organism or habitat of interest. Where trampling is relevant, the evidence and trampling intensity will be reported in the rationale. Further details. EvidenceAs a soft bodied species, Polydora ciliata is likely to be crushed and killed by an abrasive force or physical blow. However, some individuals are likely to survive as individuals can withdraw into burrows and so 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. | Intermediate | High | Low | Moderate |
Displacement [Show more]DisplacementBenchmark. Removal of the organism from the substratum and displacement from its original position onto a suitable substratum. A single event is assumed for assessment. Further details EvidencePolydora ciliata is capable of tube building throughout its life and so is able to re-establish attachment on displacement. In experimental removal of Polydora ciliata individuals of all ages which were removed from their tubes on many occasions, all 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. | Low | High | Low | Moderate |
Chemical pressures
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Intolerance | Recoverability | Sensitivity | Evidence / Confidence | |
Synthetic compound contamination [Show more]Synthetic compound contaminationSensitivity is assessed against the available evidence for the effects of contaminants on the species (or closely related species at low confidence) or community of interest. For example:
The evidence used is stated in the rationale. Where the assessment can be based on a known activity then this is stated. The tolerance to contaminants of species of interest will be included in the rationale when available; together with relevant supporting material. Further details. EvidencePolydora ciliata was abundant at polluted sites close to acidified, halogenated effluent discharge from a bromide-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. | Low | High | Low | Low |
Heavy metal contamination [Show more]Heavy metal contaminationEvidenceExperimental 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). | Intermediate | High | Low | Low |
Hydrocarbon contamination [Show more]Hydrocarbon contaminationEvidenceIn 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. This species has some brood protection which enables larvae to settle almost immediately in the nearby area (Reish, 1979). | Intermediate | High | Low | Not relevant |
Radionuclide contamination [Show more]Radionuclide contaminationEvidenceInsufficientinformation. | No information | No information | No information | Not relevant |
Changes in nutrient levels [Show more]Changes in nutrient levelsEvidencePolydora ciliata is often found in environments subject to high levels of nutrients (Sordino et al, 1989). For example, the species was abundant in areas of the Firth of Forth exposed to high levels of sewage pollution (Smyth, 1968). However, Polydora ciliata is also common in organically poor areas (Pearson & Rosenberg, 1978) and so is likely to have low intolerance to changes in nutrient concentrations. 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). | Low | High | Low | High |
Increase in salinity [Show more]Increase in salinity
EvidencePolydora 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). | Low | High | Low | High |
Decrease in salinity [Show more]Decrease in salinity
Evidence | No information | |||
Changes in oxygenation [Show more]Changes in oxygenationBenchmark. Exposure to a dissolved oxygen concentration of 2 mg/l for one week. Further details. EvidencePolydora ciliata is assessed as having low intolerance to changes in oxygenation because the species is repeatedly found at localities with oxygen deficiency (Pearson & Rosenberg, 1978). For example, in polluted harbours 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). Recovery is good because the species is able to rapidly recolonize suitable habitats. | Low | High | Low | High |
Biological pressures
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Intolerance | Recoverability | Sensitivity | Evidence / Confidence | |
Introduction of microbial pathogens/parasites [Show more]Introduction of microbial pathogens/parasitesBenchmark. Sensitivity can only be assessed relative to a known, named disease, likely to cause partial loss of a species population or community. Further details. EvidenceNo information on diseases of Polydora ciliata was found. | No information | Not relevant | No information | Not relevant |
Introduction of non-native species [Show more]Introduction of non-native speciesSensitivity assessed against the likely effect of the introduction of alien or non-native species in Britain or Ireland. Further details. EvidenceNo known non-native species compete with Polydora ciliata. | No information | No information | No information | Not relevant |
Extraction of this species [Show more]Extraction of this speciesBenchmark. Extraction removes 50% of the species or community from the area under consideration. Sensitivity will be assessed as 'intermediate'. The habitat remains intact or recovers rapidly. Any effects of the extraction process on the habitat itself are addressed under other factors, e.g. displacement, abrasion and physical disturbance, and substratum loss. Further details. EvidenceExtraction of the species is unlikely although dredging may remove populations in some habitats. Recovery is good because Polydora ciliata is iteroparous and larvae can disperse over long distances. Recolonization is rapid, usually taking place within several months of the reproductive period in the summer. | Low | High | Low | Moderate |
Extraction of other species [Show more]Extraction of other speciesBenchmark. A species that is a required host or prey for the species under consideration (and assuming that no alternative host exists) or a keystone species in a biotope is removed. Any effects of the extraction process on the habitat itself are addressed under other factors, e.g. displacement, abrasion and physical disturbance, and substratum loss. Further details. EvidenceAlthough Polydora ciliata is often associated with oysters and mussels it is not dependent on another species. | Not relevant | Not relevant | Not relevant | Not relevant |
Additional information
Importance review
Policy/legislation
- no data -
Status
National (GB) importance | - | Global red list (IUCN) category | - |
Non-native
Parameter | Data |
---|---|
Native | - |
Origin | - |
Date Arrived | - |
Importance information
- Polydora ciliata is a serious pest of oysters and mussels, but invades only the shell and does not eat the soft tissue. When infestation is heavy, the shell is weakened and this makes the mollusc more susceptible to predation by crabs. Occasionally, the inner surface of the shell is damaged by the worms and the mollusc secretes a nacreous substance to seal the perforation. The resulting blisters are an indication of attack by the species, and mussels heavily infested with the polychaete have reduced flesh content, possibly as a result of some impact on the metabolism of the mussel.
- In areas of mud tubes built by Polydora ciliata can agglomerate and form layers of mud up to an average of 20cm thick, occasionally to 50cm. These 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).
Bibliography
Bryan, G.W., 1984. Pollution due to heavy metals and their compounds. In Marine Ecology: A Comprehensive, Integrated Treatise on Life in the Oceans and Coastal Waters, vol. 5. Ocean Management, part 3, (ed. O. Kinne), pp.1289-1431. New York: John Wiley & Sons.
Crisp, D.J. (ed.), 1964. The effects of the severe winter of 1962-63 on marine life in Britain. Journal of Animal Ecology, 33, 165-210.
Daro, M.H. & Polk, P., 1973. The autecology of Polydora ciliata along the Belgian coast. Netherlands Journal of Sea Research, 6, 130-140.
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.
Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.
Grassle, J.F. & Grassle, J.P., 1974. Opportunistic life histories and genetic systems in marine benthic polychaetes. Journal of Marine Research, 32, 253-284.
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.
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.
Gulliksen, B., 1977. Studies from the “UWL Helgoland” on the macrobenthic fauna of rocks and boulders in Lübeck Bay (western Baltic Sea). Helgolander Wissenschaftliche Meeresuntersuchungen, 30(1-4), 519-526.
Harms, J. & Anger, K., 1983. Seasonal, annual, and spatial variation in the development of hard bottom communities. Helgoländer Meeresuntersuchungen, 36, 137-150.
Hayward, P.J. & Ryland, J.S. (ed.) 1995b. Handbook of the marine fauna of North-West Europe. Oxford: Oxford University Press.
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.
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
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.
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.
Murina, V., 1997. Pelagic larvae of Black Sea Polychaeta. Bulletin of Marine Science, 60, 427-432.
Mustaquim, J., 1986. Morphological variation in Polydora ciliata complex (Polychaeta, Annelida). Zoological Journal of the Linnean Society, 86, 75-88.
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.
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.
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.
Rosenberg, R., 1977. Benthic macrofaunal dynamics, production, and dispersion in an oxygen-deficient estuary of west Sweden. Journal of Experimental Marine Biology and Ecology, 26, 107-33.
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.
Smyth, J.C., 1968. The fauna of a polluted site in the Firth of Forth. Helgolander Wissenschaftliche Meeresuntersuchungen, 17, 216-233.
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.
Datasets
Bristol Regional Environmental Records Centre, 2017. BRERC species records recorded over 15 years ago. Occurrence dataset: https://doi.org/10.15468/h1ln5p accessed via GBIF.org on 2018-09-25.
Centre for Environmental Data and Recording, 2018. IBIS Project Data. Occurrence dataset: https://www.nmni.com/CEDaR/CEDaR-Centre-for-Environmental-Data-and-Recording.aspx accessed via NBNAtlas.org on 2018-09-25.
National Trust, 2017. National Trust Species Records. Occurrence dataset: https://doi.org/10.15468/opc6g1 accessed via GBIF.org on 2018-10-01.
NBN (National Biodiversity Network) Atlas. Available from: https://www.nbnatlas.org.
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Last Updated: 24/04/2007