Polydora sp. tubes on moderately exposed sublittoral soft rock

Summary

UK and Ireland classification

Description

Large patches of chalk and soft limestone are occasionally covered entirely by Polydora sp. tubes to the exclusion of almost all other species. This tends to occur in highly turbid conditions and spans the infralittoral and circalittoral in limestone areas such as the Great and Little Ormes (North Wales) and Gower (South Wales). It is even present on the lower shore in the Severn estuary. The boring form of the sponge Cliona celata often riddles the surface layer of the stone. Other sponges present include Halichondria panicea, Haliclona oculata and Hymeniacidon perlevis (syn. Hymeniacidon perleve). Polydora sp. also frequently occurs in small patches as part of other biotopes (e.g. FluCoAs). Other species present include Alcyonium digitatum, Sarcodictyon roseum, the hydroids Halecium halecinum, Abietinaria abietina and Tubularia indivisa, the ascidians Clavelina lepadiformis, Botryllus schlosseri and Morchellium argus, the anemones Urticina felina, Metridium senile (syn. Metridium senile) and Cylista elegans and the bryozoans Flustra foliacea and a crisiid turf. The starfish Asterias rubens, the crabs Inachus phalangium and Carcinus maenas, the polychaete Spirobranchus triqueter (syn. Spirobranchus triqueter), the barnacle Balanus crenatus and the brittlestar Ophiothrix fragilis may also be seen. Please note: this biotope may extend into the infralittoral and littoral zone in areas where water turbidity is sufficiently high. (Information taken from Connor et al., 2004).

Depth range

Lower shore, 0-5 m, 5-10 m, 10-20 m

Additional information

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Habitat review

Ecology

Ecological and functional relationships

  • In areas of mud, the tubes built by Polydora ciliata can agglomerate and form layers of mud up to an average of 20 cm 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).
  • 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 about 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).
  • Polydora ciliata is predated upon by urchins and in Helgoland there is a close relationship between the distribution of Polydora ciliata and Echinus esculentus. Echinus esculentus grazes almost exclusively on the Polydora ciliata carpets and takes its main food not from biodetritus and animals living between the Polydora chimneys but by feeding on the worm itself. To reach the worm, Echinus esculentus has to scrape away between 0.5and 1.2 cm of solid rock and this feeding behaviour is responsible for the bioerosion of rocks in the Helgoland area by an estimated 1cm per annum (Krumbein & Van der Pers, 1974).

Seasonal and longer term change

The early reproductive period of Polydora ciliata often enables the species to be the first 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 hard substrata are covered with the thickest layer of 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 cannot 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 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 form layers of muddy tubes on soft rock. 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 MCR.Pol is mostly secondary, derived from detritus and organic material. 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, 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 substrate 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 substrates and the larvae of the species settle preferentially on substrates 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

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Preferences & Distribution

Habitat preferences

Depth Range Lower shore, 0-5 m, 5-10 m, 10-20 m
Water clarity preferences
Limiting Nutrients Not relevant
Salinity preferences Full (30-40 psu)
Physiographic preferences Open coast
Biological zone preferences Circalittoral
Substratum/habitat preferences Bedrock
Tidal strength preferences Moderately strong 1 to 3 knots (0.5-1.5 m/sec.), Weak < 1 knot (<0.5 m/sec.)
Wave exposure preferences Moderately exposed
Other preferences Soft rock including chalk, limestone and sandstone

Additional Information

Species composition

Species found especially in this biotope

    Rare or scarce species associated with this biotope

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    Additional information

    Sensitivity review

    Sensitivity characteristics of the habitat and relevant characteristic species

    CR.MCR.SfR.Pol is a sublittoral biotope occurring in moderately exposed areas with strong, but also moderately strong and weak tidal streams (Connor et al., 2004). This biotope is defined by occurring in soft rock, such as chalk and soft limestone, in areas where water turbidity is high. Large patches of chalk and soft limestone are occasionally covered entirely by Polydora sp. tubes to the exclusion of almost all other species (Daro & Polk, 1973). The species readily overgrows other species with a flat morphology and feeds by scraping its palps about its tubes, which would inhibit the development of settling larvae of other species. For this reason, the mat of Polydora spp. tubes is considered as the defining characteristic of this biotope. In the north west of Europe, Polydora ciliata has been mainly associated to limestone rock and stones (Hayward & Ryland, 1995b), so it is likely that this species is characteristic of this this biotope and is therefore the focus of this assessment as an example of tube building Polydora spp. In this biotope Polydora spp. (and their tubes) are therefore the key structuring and defining element. A range of other epifaunal species are present in patches including sponges, hydroids, barnacles, ascidians and anemones. These contribute to species richness and diversity but are not considered key characterizing, defining or structuring species and are not considered within the assessments. More information on these species can be found in other biotope assessments available on this website. 

    The soft rock substratum is considered a key element defining the habitat and supporting the development of this biotope. The substratum is therefore considered in sensitivity assessments where the pressure may alter or change it.

    Resilience and recovery rates of habitat

    Polyodra is a small, sedentary, burrowing polychaete worm up to 3 cm long. All Polydora spp. make a U-shaped tube from small particles (Hayward & Ryland, 1995b). Polydora ciliata usually burrows into substrata containing calcium carbonate such as limestone, chalk and clay, as well as shells or oysters, mussels and periwinkles (Fish & Fish, 1996). The sexes are separate and breeding has been recorded in spring in a number of locations. In northern England, spawning has been recorded to occur from February until June and three or four generations succeed one another during the spawning period (Gudmundsson, 1985). Eggs are laid in a string of capsules that are attached by two threads to the wall of the burrow (Fish & Fish, 1996). After a week the larvae emerge and are believed to have a pelagic life of from 2-6 weeks before settling. Length of life is no more than 1 year (Fish & Fish, 1996). Larvae of Polydora ciliata have been collected as far as 118 km offshore (Murina, 1997). Larvae settle on specific substratum types, selecting rocks according to their physical properties or sediment depending on substrate particle size. Adults of Polydora ciliata produce a 'mud' resulting from the perforation of soft rock substrates and the larvae of the species settle preferentially on substrates covered with mud (Lagadeuc, 1991). The tubes built by Polydora agglomerate sometimes to form layers of mud up to an average of 20 cm thick. However, it may take several years for a Polydora ciliata 'mat' to reach a significant size (Hill, 2007). However, interspecific competition and heavy mortality of the larvae have been observed on Polydora mats (Daro & Polk, 1973).

    The early reproductive period of Polydora ciliata often enables the species to be the first 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. 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 cannot 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 in June. Recolonization of the habitat later in the year may be inhibited by other species that colonize and compete for rock and therefore later settlements may not result in the formation of this biotope.

    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.

    Resilience assessment: Removal of the characterizing species Polydora would likely result in the biotope being lost and re-classified. Polydora is short lived species, reaching maturity within a few months, and communities appear to be annual, sustained by three or even four generations succeeding one another and resulting in planktonic larvae that can be found throughout the year (Daro & Polk, 1973). In the event of a large portion of the Polydora community being lost, the biotope is likely to reach maturity very. For as long as the substrate nature of the biotope remains suitable for the settlement of Polydora recruits, the biotope is likely to recover fully within two years, hence resilience has been assessed as High.

    NB: The resilience and the ability to recover from human induced pressures is a combination of the environmental conditions of the site, the frequency (repeated disturbances versus a one-off event) and the intensity of the disturbance. Recovery of impacted populations will always be mediated by stochastic events and processes acting over different scales including, but not limited to, local habitat conditions, further impacts and processes such as larval-supply and recruitment between populations. Full recovery is defined as the return to the state of the habitat that existed prior to impact.  This does not necessarily mean that every component species has returned to its prior condition, abundance or extent but that the relevant functional components are present and the habitat is structurally and functionally recognizable as the initial habitat of interest. It should be noted that the recovery rates are only indicative of the recovery potential.

    Hydrological Pressures

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    ResistanceResilienceSensitivity
    Temperature increase (local) [Show more]

    Temperature increase (local)

    Benchmark. A 5°C increase in temperature for one month, or 2°C for one year. Further detail

    Evidence

    Murina (1997) categorized 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, where sea temperatures varied between 0.5 and 17°C (Dorsett, 1961). Growth rates may increase if temperature rises. For example, at Whitstable in Kent, Dorsett (1961) found that a rapid increase in growth of Polydora ciliata coincided with the rising temperature of the seawater during March.

    Sensitivity assessment: Typical surface water temperatures around the UK coast vary, seasonally from 4-19°C (Huthnance, 2010). No information was found on the maximum temperature tolerated by Polydora ciliata. However, it is likely that the species is able to resist a long-term increase in temperature of 2°C and may resist a short-term increase of 5°C. Resistance and resilience are therefore assessed as High and the biotope is judged as Not Sensitive.

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    High
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    Not sensitive
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    Temperature decrease (local) [Show more]

    Temperature decrease (local)

    Benchmark. A 5°C decrease in temperature for one month, or 2°C for one year. Further detail

    Evidence

    Murina (1997) categorized 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). During the extremely cold winter of 1962/63 Polydora ciliata was apparently unaffected, when temperature anomalies of between 2.5-5.8°C where observed (Crisp, 1964).

    Sensitivity assessment: Typical surface water temperatures around the UK coast vary, seasonally from 4-19°C (Huthnance, 2010). Polydora ciliata is likely to be able to resist a long-term decrease in temperature of 2°C and may resist a short-term decrease of 5°C.  Temperature may act as a spawning cue and an acute or chronic decrease may result in some delay in spawning, however this is not considered to impact the adult population and may be compensated by later spawning events. Resistance and resilience are therefore assessed as High and the biotope judged as Not Sensitive.

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    High
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    Not sensitive
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    Salinity increase (local) [Show more]

    Salinity increase (local)

    Benchmark. A increase in one MNCR salinity category above the usual range of the biotope or habitat. Further detail

    Evidence

    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.

    Sensitivity assessment: A long-term increase in salinity at the pressure benchmark level is likely to result in the death of many individuals. Resistance is therefore assessed as Low and resilience is likely to be High so the biotope is considered to have Low sensitivity to an increase in salinity at the pressure benchmark level.

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    High
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    Low
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    Salinity decrease (local) [Show more]

    Salinity decrease (local)

    Benchmark. A decrease in one MNCR salinity category above the usual range of the biotope or habitat. Further detail

    Evidence

    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).

    Sensitivity assessment: Records indicate CR.MCR.SfR.Pol occurs in areas of variable (18-35 ppt) salinity (Connor et al., 2004). Polydora ciliata is therefore likely to resist a decrease in salinity at the pressure benchmark level. Resistance is therefore assessed as High and resilience as High (by default) and the biotope is considered Not Sensitive to a decrease in salinity at the pressure benchmark level.

    High
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    High
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    Not sensitive
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    Water flow (tidal current) changes (local) [Show more]

    Water flow (tidal current) changes (local)

    Benchmark. A change in peak mean spring bed flow velocity of between 0.1 m/s to 0.2 m/s for more than one year. Further detail

    Evidence

    CR.MCR.SfR.Pol is recorded in a range of tidal streams, including strong (1.5-3 m/s), moderately strong (0.5-1.5 m/s) and weak tidal stream conditions (<0.5 m/s) (Connor et al., 2004).

    Polydora ciliata colonized test panels in Helgoland in three areas, two exposed to strong tidal currents and one site sheltered from currents (Harms & Anger, 1983). Very strong water flows may sweep away Polydora colonies, where these are present as a thick layer of mud on a hard substratum.

    The most damaging effect of increased flow rate (above the pressure benchmark) could be the erosion of the soft rock substratum as this could eventually lead to loss of the habitat.

    Sensitivity assessment: CR.MCR.SfR.Pol is recorded in a range of tidal streams, including strong, moderately strong and weak tidal stream conditions (Connor et al., 2004). A change in water flow at the benchmark level of 0.1-0.2 m/s is therefore likely to fall within the normal range of water flows experienced by the biotope. Resistance and resilience are therefore considered to be High and the biotope is assessed as Not Sensitive to a change in water flow rate at the pressure benchmark level.

    High
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    High
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    Not sensitive
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    Emergence regime changes [Show more]

    Emergence regime changes

    Benchmark.  1) A change in the time covered or not covered by the sea for a period of ≥1 year or 2) an increase in relative sea level or decrease in high water level for ≥1 year. Further detail

    Evidence

    Changes in emergence are Not Relevant to the biotope, which is restricted to fully subtidal/circalittoral conditions. The pressure benchmark is relevant only to littoral and shallow sublittoral fringe biotopes.

    Not relevant (NR)
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    Not relevant (NR)
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    Not relevant (NR)
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    Wave exposure changes (local) [Show more]

    Wave exposure changes (local)

    Benchmark. A change in near shore significant wave height of >3% but <5% for more than one year. Further detail

    Evidence

    The biotope is found in moderately exposed sites (Connor et al., 2004). Feeding of Polydora ciliata may be impaired in strong wave action and changes in wave exposure may also influence the supply of particulate matter for tube building activities. Decreases in wave exposure may influence the supply of particulate matter 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.

    Potentially the most damaging effect of increased wave heights would be the erosion of the soft rock substratum as this could eventually lead to loss of the habitat.

    Sensitivity assessment. Some erosion will occur naturally and storm events may be more significant in loss and damage of soft rocks than changes in wave height at the pressure benchmark. The biotope is therefore considered to have High resistance to changes at the pressure benchmark where these do not lead to increased erosion of the substratum. Resilience is therefore assessed as High and the biotope is considered to be Not Sensitive, at the pressure benchmark.

    High
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    High
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    Not sensitive
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    Chemical Pressures

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    ResistanceResilienceSensitivity
    Transition elements & organo-metal contamination [Show more]

    Transition elements & organo-metal contamination

    Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

    Evidence

    This pressure is Not assessed but evidence is presented where available.

    Experimental studies with various species suggest that polychaete worms are quite resistant 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).

    Not Assessed (NA)
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    Not assessed (NA)
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    Not assessed (NA)
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    Hydrocarbon & PAH contamination [Show more]

    Hydrocarbon & PAH contamination

    Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

    Evidence

    This pressure is Not assessed but evidence is presented where available.

    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 the 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).

    Not Assessed (NA)
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    Not assessed (NA)
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    Synthetic compound contamination [Show more]

    Synthetic compound contamination

    Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

    Evidence

    This pressure is Not assessed but evidence is presented where available.

    Polydora 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 resistant of distilling and petrochemical industrial waste in Scotland.

    Not Assessed (NA)
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    Not assessed (NA)
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    Not assessed (NA)
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    Radionuclide contamination [Show more]

    Radionuclide contamination

    Benchmark. An increase in 10µGy/h above background levels. Further detail

    Evidence

    No evidence was found

    No evidence (NEv)
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    Not relevant (NR)
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    No evidence (NEv)
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    Introduction of other substances [Show more]

    Introduction of other substances

    Benchmark. Exposure of marine species or habitat to one or more relevant contaminants via uncontrolled releases or incidental spills. Further detail

    Evidence

    This pressure is Not assessed.

    Not Assessed (NA)
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    Not assessed (NA)
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    De-oxygenation [Show more]

    De-oxygenation

    Benchmark. Exposure to dissolved oxygen concentration of less than or equal to 2 mg/l for one week (a change from WFD poor status to bad status). Further detail

    Evidence

    Polydora ciliata is frequently 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). Furthermore, in a study investigating a polychaete community in the north west Black Sea, Polydora ciliata was observed in all four study sites, including those severely affected by eutrophication and hypoxia as a result of discharges of wastewaters (Vorobyova et al., 2008). However, Polydora ciliata is unlikely to be able to resist anoxic conditions. Hansen et al. (2002) reported near total extinction of all metazoan in the Mariager Fjord (Denmark), including Polydora spp. after a severe hypoxia event that resulted in complete anoxia in the water column for two weeks. Additionally, Como & Magni (2009) investigated seasonal variations in benthic communities known to be affected by episodic events of hypoxia. The authors observed that abundance of Polydora ciliata varied seasonally, decreasing during the summer months, and suggested it could be explained by the occurrence of hypoxic/anoxic conditions and sulphidic sediments during the summer. No details of the levels of dissolved oxygen leading to these community responses were provided.

    Sensitivity assessment: Polydora ciliata is frequently found at localities with oxygen deficiency (Pearson & Rosenberg, 1978) and seems to only be affected by severe de-oxygenation episodes. Resistance to de-oxygenation at the pressure benchmark level is likely to be High. Opportunistic Polydora spp. have also repeatedly been reported amongst the first to recover hypoxia events (Hansen et al., 2002; Van Colen et al., 2010) so resilience is likely to also be High. The biotope is therefore considered Not Sensitive to exposure to dissolved oxygen concentration of less than or equal to 2 mg/l for 1 week.

    High
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    High
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    Not sensitive
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    Nutrient enrichment [Show more]

    Nutrient enrichment

    Benchmark. Compliance with WFD criteria for good status. Further detail

    Evidence

    Polydora ciliata is often found in environments subject to high levels of nutrient input. For example, the species was abundant in areas of the Firth of Forth exposed to high levels of sewage pollution (Smyth, 1968), 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). The extensive growths of Polydora ciliata in mat formations were recorded at West Ganton, in the Firth of Forth, prior to the introduction of the Sewage Scheme (Read et al., 1983). The abundance of the species was probably associated with their ability to use the increased availability of nutrients as a food source and silt for tube building.

    Sensitivity assessment: The characterizing species of this biotope is likely to be able to resist nutrient enrichment. The biotope is considered Not Sensitive at the pressure benchmark that assumes compliance with good status as defined by the WFD.

    Not relevant (NR)
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    Not relevant (NR)
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    Not sensitive
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    Organic enrichment [Show more]

    Organic enrichment

    Benchmark. A deposit of 100 gC/m2/yr. Further detail

    Evidence

    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). However, Callier et al. (2007) investigated the spatial distribution of macrobenthos under a suspended mussel culture, in eastern Canada, where the sedimentation of organic matter to the bottom was approx. 1-3 gC/m2/d. Polydora ciliata was recorded as absent in the sites under the suspended mussel farm after one year and as dominant in reference areas of the study.  It should be noted that the organic matter input from the mussel farm exceeds the pressure benchmark.

    Como & Magni (2009) investigated seasonal variations in benthic communities known to be affected by episodic events of sediment over-enrichment. The authors observed that abundance of Polydora ciliata varied seasonally, and suggested this could be a result of major accumulation of organic C-bounding fine sediments in the study site.

    Studies by Almeda et al. (2009) and Pedersen et al. (2010) investigated larval energetic requirements for Polydora ciliata, and suggested maximum growth rates were reached at food concentrations ranging from 2.5 to 1.4 μg C/ml depending on larval size, energetic carbon requirements of 0.09 to 3.15 μg C l/d, respectively.

    Polydora ciliata can also occur in organically poor areas (Pearson & Rosenberg, 1978).

    Borja et al. (2000) and Gittenberger & van Loon (2011) in the development of an AMBI index to assess disturbance (including organic enrichment) both assigned Polydora ciliata to their Ecological Group IV ‘Second-order opportunistic species present in slight to pronounced unbalanced situations’.

    Sensitivity assessment: The evidence presented suggests Polydora ciliata may not be able to resist organic enrichment that exceeds the pressure benchmark level (Callier et al., 2007) but may not be affected by at the benchmark level. Resistance and resilience are therefore assessed as High and the biotope is considered Not Sensitive to organic enrichment (deposit of 100 gC/m2/yr).

    High
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    High
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    Not sensitive
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    Physical Pressures

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    ResistanceResilienceSensitivity
    Physical loss (to land or freshwater habitat) [Show more]

    Physical loss (to land or freshwater habitat)

    Benchmark. A permanent loss of existing saline habitat within the site. Further detail

    Evidence

    All marine habitats and benthic species are considered to have a resistance of None to this pressure and to be unable to recover from a permanent loss of habitat (Resilience is Very Low). Sensitivity within the direct spatial footprint of this pressure is therefore High. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure.

    None
    High
    High
    High
    Help
    Very Low
    High
    High
    High
    Help
    High
    High
    High
    High
    Help
    Physical change (to another seabed type) [Show more]

    Physical change (to another seabed type)

    Benchmark. Permanent change from sedimentary or soft rock substrata to hard rock or artificial substrata or vice-versa. Further detail

    Evidence

    CR.MCR.SfR.Pol is characterized by the soft rock substratum which supports populations of Polydora sp. tubes. A change to a sedimentary, hard rock or artificial substratum would result in the loss of Polydora, significantly altering the character of the biotope. The biotope would be lost and/or re-classified.

    Sensitivity assessment: Resistance to the pressure is considered None, and resilience Very Low based on the loss of suitable substratum to support the community of the characterizing species of Polydora. Sensitivity has been assessed as High. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure. 

    None
    High
    High
    High
    Help
    Very Low
    High
    High
    High
    Help
    High
    High
    High
    High
    Help
    Physical change (to another sediment type) [Show more]

    Physical change (to another sediment type)

    Benchmark. Permanent change in one Folk class (based on UK SeaMap simplified classification). Further detail

    Evidence

    Not Relevant to biotopes occurring on bedrock.

    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Habitat structure changes - removal of substratum (extraction) [Show more]

    Habitat structure changes - removal of substratum (extraction)

    Benchmark. The extraction of substratum to 30 cm (where substratum includes sediments and soft rock but excludes hard bedrock). Further detail

    Evidence

    Removal of the substratum to 30 cm would result in the loss of Polydora sp. tubes.  Resistance to the pressure is considered None, and resilience Very Low based on the loss of suitable substratum to support the community of the characterizing species of Polydora. Sensitivity has been assessed as High. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure.

    None
    High
    High
    High
    Help
    Very Low
    High
    High
    High
    Help
    High
    High
    High
    High
    Help
    Abrasion / disturbance of the surface of the substratum or seabed [Show more]

    Abrasion / disturbance of the surface of the substratum or seabed

    Benchmark. Damage to surface features (e.g. species and physical structures within the habitat). Further detail

    Evidence

    This biotope is characterized by epifauna occurring on hard rock substratum. The tubes of Polydora spp. are likely to be removed by abrasion as these project above the surface and are not physically robust. Other epifauna associated with this biotope are also likely to be damaged and/or removed by surface abrasion. Some species such as anemones and sponges may be able to rapidly repair damage while others may recolonize rapidly, e.g. barnacles.  

    Sensitivity assessment.  The characterizing Polydora community in this biotope, is considered likely to be damaged and removed by abrasion. As a soft bodied species, Polydora ciliata is likely to be crushed and killed by an abrasive force or physical blow. Erect epifauna are directly exposed to this pressure which would displace, damage and remove individuals. Resistance to abrasion is considered None. However, Polydora is likely to be able to re-establish the lost community rapidly, so resilience of the biotope is assessed as High with the biotope considered to have Medium sensitivity to abrasion or disturbance of the surface of the seabed. The substratum is unable to recover from damage and therefore the biotope would be considered highly sensitivity to abrasion that damaged or removed the soft rock substratum.

    None
    High
    High
    High
    Help
    High
    High
    Medium
    High
    Help
    Medium
    High
    Medium
    High
    Help
    Penetration or disturbance of the substratum subsurface [Show more]

    Penetration or disturbance of the substratum subsurface

    Benchmark. Damage to sub-surface features (e.g. species and physical structures within the habitat). Further detail

    Evidence

    Activities that disturb the surface of the mat and penetrate below the surface would remove a significant proportion of the Polydora tubes within the direct area of impact. Biotope resistance is therefore assessed as None and recovery is assessed as High based on the assumption that the suitable substratum to support the community of the characterizing species of Polydora would only be damaged, not lost. Sensitivity is therefore assessed as Medium. The substratum is unable to recover from damage and therefore the biotope would be considered highly sensitivity to physical disturbance that damaged or removed the soft rock substratum. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure.

    None
    High
    High
    High
    Help
    High
    High
    High
    High
    Help
    Medium
    High
    High
    High
    Help
    Changes in suspended solids (water clarity) [Show more]

    Changes in suspended solids (water clarity)

    Benchmark. A change in one rank on the WFD (Water Framework Directive) scale e.g. from clear to intermediate for one year. Further detail

    Evidence

    CR.MCR.SfR.Pol tends to occur in highly turbid areas (Connor et al., 2004). In the Firth of Forth, Polydora ciliata formed extensive mats in areas that had an average of 68 mg/l suspended solids and a maximum of approximately 680 mg/l, indicating the species is able to tolerate different levels 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).

    Suspended sediment and siltation of particles is important for tube building in Polydora ciliata so a decrease in suspended solids may reduce tube building or the thickness of the mud surrounding the 'colonies'. Daro & Polk (1973) reported that the success of Polydora is directly related to the quantities of muds of any origin carried along by rivers or coastal currents.

    An increase in turbidity, reducing light availability may reduce primary production by phytoplankton in the water column. A reduction in primary production in the water column may result indirectly in reduced food supply to the detritus feeding Polydora which in turn may affect growth rates and fecundity.

    Sensitivity assessment: An increase in suspended solids at the pressure benchmark level is unlikely to affect the characterizing species of this biotope. However, a decrease in suspended matter in the biotope could result in limitation of material for tube building activity of Polydora and also in the substrate no longer being suitable for colonization by new recruits. Resistance of the biotope is therefore considered to be Low (loss of 25-75%) and resilience is High (following a return to normal conditions) so the biotope is considered to have Low sensitivity to a decrease in suspended solids at the pressure benchmark level.

    Low
    High
    High
    High
    Help
    High
    High
    Medium
    High
    Help
    Low
    High
    Medium
    High
    Help
    Smothering and siltation rate changes (light) [Show more]

    Smothering and siltation rate changes (light)

    Benchmark. ‘Light’ deposition of up to 5 cm of fine material added to the seabed in a single discrete event. Further detail

    Evidence

    Adults of Polydora ciliata produce a 'mud' resulting from the perforation of soft rock substrates (Lagadeuc, 1991). A Polydora mud can be up to 50 cm thick, but the animals themselves occupy only the first few centimetres. They either elongate their tubes to reach the surface, or leave them to rebuild close to the surface.

    Munari & Mistri (2014) investigated the spatio-temporal variation pattern of a benthic community following deposition of dredged material, at a maximum thickness of 30–40 cm. Polydora ciliata was amongst the first colonizers of the newly deposited sediments. The authors suggested that it was possible that the individuals migrated vertically through the deep layer of dredged sand. This was based on the results of Roberts et al. (1998) who suggested 15 cm as the maximum depth of overburden through which benthic infauna can successfully migrate. After one year, no adverse impact of sand disposal on the benthic fauna was detected on the study site.

    Sensitivity assessment: Based on the evidence presented by Munari & Mistri (2014), Polydora ciliata is considered likely to resist smothering by 5 cm of sediment. Resistance and resilience are therefore assessed as High and the biotope is considered Not Sensitive to a ‘light’ deposition of up to 5 cm of fine material in a single discrete event.

    High
    High
    Medium
    High
    Help
    High
    High
    High
    High
    Help
    Not sensitive
    High
    Medium
    High
    Help
    Smothering and siltation rate changes (heavy) [Show more]

    Smothering and siltation rate changes (heavy)

    Benchmark. ‘Heavy’ deposition of up to 30 cm of fine material added to the seabed in a single discrete event. Further detail

    Evidence

    Adults of Polydora ciliata produce a 'mud' resulting from the perforation of soft rock substrates (Lagadeuc, 1991). A Polydora mud can be up to 50 cm thick, but the animals themselves occupy only the first few centimetres. They either elongate their tubes, or leave them to rebuild close to the surface.

    Munari & Mistri (2014) investigated the spatio-temporal variation pattern of a benthic community following deposition of dredged material, at a maximum thickness of 30–40 cm. Polydora ciliata was amongst the first colonizers of the newly deposited sediments. The authors suggested that it was possible that the individuals migrated vertically through the deep layer of dredged sand. This was based on the results of Roberts et al. (1998) who suggested 15 cm as the maximum depth of overburden through which benthic infauna can successfully migrate. After one year, no adverse impact of sand disposal on the benthic fauna was detected on the study site.

    Sensitivity assessment: Polychaete species have been reported to migrate through depositions of sediment greater that the benchmark (30 cm of fine material added to the seabed in a single discrete event) (Maurer et al., 1982). However, it is not clear whether Polydora ciliata is likely to be able to migrate through a maximum thickness of fine sediment that would compare to that investigated by Munari & Magni (2014) because muds tend to be more cohesive and compacted than sand. Some mortality of the characterizing species is likely to occur. Resistance is therefore assessed as Low and resilience as High and the biotope is considered to have Low sensitivity to a ‘heavy’ deposition of up to 30 cm of fine material in a single discrete event.

    Low
    High
    High
    Medium
    Help
    High
    High
    Medium
    High
    Help
    Low
    High
    Medium
    Medium
    Help
    Litter [Show more]

    Litter

    Benchmark. The introduction of man-made objects able to cause physical harm (surface, water column, seafloor or strandline). Further detail

    Evidence

    Not assessed.

    Not Assessed (NA)
    NR
    NR
    NR
    Help
    Not assessed (NA)
    NR
    NR
    NR
    Help
    Not assessed (NA)
    NR
    NR
    NR
    Help
    Electromagnetic changes [Show more]

    Electromagnetic changes

    Benchmark. A local electric field of 1 V/m or a local magnetic field of 10 µT. Further detail

    Evidence

    No Evidence is available on which to assess this pressure.

    No evidence (NEv)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    No evidence (NEv)
    NR
    NR
    NR
    Help
    Underwater noise changes [Show more]

    Underwater noise changes

    Benchmark. MSFD indicator levels (SEL or peak SPL) exceeded for 20% of days in a calendar year. Further detail

    Evidence

    Polydora ciliata may respond to vibrations from predators or bait diggers by retracting their palps into their tubes. However, the species is unlikely to be affected by noise pollution and so the biotope is assessed as Not Sensitive.

    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Introduction of light or shading [Show more]

    Introduction of light or shading

    Benchmark. A change in incident light via anthropogenic means. Further detail

    Evidence

    CR.MCR.SfR.Pol is a circalittoral biotope (Connor et al., 2004) and therefore, not directly dependent on sunlight. 

    Sensitivity assessment. Although Polydora spp. can perceive light, this pressure is not considered relevant. The biotope is considered to have High resistance and, by default, High resilience and therefore is Not Sensitive to the introduction of light.

    High
    Low
    NR
    NR
    Help
    High
    High
    High
    High
    Help
    Not sensitive
    Low
    NR
    NR
    Help
    Barrier to species movement [Show more]

    Barrier to species movement

    Benchmark. A permanent or temporary barrier to species movement over ≥50% of water body width or a 10% change in tidal excursion. Further detail

    Evidence

    Not Relevant to biotopes restricted to open waters.

    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Death or injury by collision [Show more]

    Death or injury by collision

    Benchmark. Injury or mortality from collisions of biota with both static or moving structures due to 0.1% of tidal volume on an average tide, passing through an artificial structure. Further detail

    Evidence

    Not Relevant to seabed habitats. NB. Collision by grounding vessels is addressed under surface abrasion.

    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Visual disturbance [Show more]

    Visual disturbance

    Benchmark. The daily duration of transient visual cues exceeds 10% of the period of site occupancy by the feature. Further detail

    Evidence

    Polydora ciliata exhibits shadow responses and withdraws its palps into its burrow, which is believed to be a defence against predation. The withdrawal of the palps interrupts feeding and possibly respiration, although the species also shows habituation of the response (Kinne, 1970).  Polydora is unlikely to be sensitive to visual disturbance caused by passing shipping but may respond to passing divers at close range. Resistance and resilience are, therefore, assessed as 'High' and the biotope judged as 'Not Sensitive' to visual disturbance.

    High
    Low
    Low
    Low
    Help
    High
    High
    High
    High
    Help
    Not sensitive
    Low
    Low
    Low
    Help

    Biological Pressures

    Use [show more] / [show less] to open/close text displayed

    ResistanceResilienceSensitivity
    Genetic modification & translocation of indigenous species [Show more]

    Genetic modification & translocation of indigenous species

    Benchmark. Translocation of indigenous species or the introduction of genetically modified or genetically different populations of indigenous species that may result in changes in the genetic structure of local populations, hybridization, or change in community structure. Further detail

    Evidence

    The key characterizing species in the biotope are not cultivated or likely to be translocated. This pressure is therefore considered Not relevant.

    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Introduction or spread of invasive non-indigenous species [Show more]

    Introduction or spread of invasive non-indigenous species

    Benchmark. The introduction of one or more invasive non-indigenous species (INIS). Further detail

    Evidence

    Crepidula fornicata larvae require hard substrata for settlement. It prefers muddy gravelly, shell-rich, substrata that include gravel, or shells of other Crepidula, or other species e.g., oysters, and mussels. It is highly gregarious and seeks out adult shells for settlement, forming characteristic ‘stacks’ of adults. But it also recorded from rock, artificial substrata, and Sabellaria alveolata reefs (Blanchard, 1997, 2009; Bohn et al., 2012, 2013a, 2013b, 2015; De Montaudouin et al., 2018; Hinz et al., 2011; Helmer et al., 2019; Powell-Jennings & Calloway, 2018; Preston et al., 2020; Tillin et al., 2020). Close examination of the literature (2023) shows that evidence of its colonization and density on bedrock in the infralittoral or circalittoral was lacking. Tillin et al. (2020) suggested that Crepidula could colonize circalittoral rock due to its presence on tide-swept rough grounds in the English Channel (Hinz et al., 2011). However, Hinz et al. (2011) reported that Crepidula fornicata only dominated one assemblage (with an average of 181 individuals per trawl) on gravel substratum with boulders. Bohn et al. (2015) noted that Crepidula occurred at low density or was absent in areas dominated by boulders, and Bohn et al. (2013a, 2013b, 2015) and Preston et al. (2020) showed that while Crepidula could settle on slate panels or ‘stone’ it preferred shell, especially that of conspecifics. In addition, no evidence was found of the effect of Crepidula populations on faunal turf-dominated habitats. It was only recorded at low density (0.1-0.9/m2) in one faunal turf biotope (CR.MCR.CFaVS.CuSpH.As) (JNCC, 2015). Faunal turfs are dominated by suspension feeders so larval predation is probably high, which may prevent colonization by Crepidula. Also, faunal turf species actively compete for space and many are fast growing and opportunistic, so may out-compete Crepidula for space even if it gained a foothold in the community. 

    Sensitivity assessment. The circalittoral rock characterizing this biotope is likely to be unsuitable for the colonization by Crepidula fornicata due to the moderately wave exposed conditions, in which wave action and storms may mitigate or prevent the colonization by Crepidula at high densities, although Crepidula has been recorded from areas of strong tidal streams (Hinz et al., 2011). In addition, no evidence was found of the effect of Crepidula populations on faunal turf-dominated habitats or infralittoral or circalittoral rock habitats. At present, there is 'Insufficient evidence' to suggest that the circalittoral rock biotopes are sensitive to colonization by Crepidula fornicata or other invasive species; further evidence is required. 

    Insufficient evidence (IEv)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Insufficient evidence (IEv)
    NR
    NR
    NR
    Help
    Introduction of microbial pathogens [Show more]

    Introduction of microbial pathogens

    Benchmark. The introduction of relevant microbial pathogens or metazoan disease vectors to an area where they are currently not present (e.g. Martelia refringens and Bonamia, Avian influenza virus, viral Haemorrhagic Septicaemia virus). Further detail

    Evidence

    Introduced organisms (especially parasites or pathogens) are a potential threat in all coastal ecosystems. However, so far, no information was found on microbial pathogens affecting Polydora ciliata.

    Sensitivity assessment. The biotope is judged to have High resistance to this pressure. By default resilience is assessed as High and the biotope is classed as Not Sensitive.

    High
    Low
    NR
    NR
    Help
    High
    High
    High
    High
    Help
    Not sensitive
    Low
    NR
    NR
    Help
    Removal of target species [Show more]

    Removal of target species

    Benchmark. Removal of species targeted by fishery, shellfishery or harvesting at a commercial or recreational scale. Further detail

    Evidence

    CR.MCR.SfR.Pol is currently not targeted by commercial fisheries and hence not directly affected by this pressure. This pressure is therefore considered Not Relevant.

    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Not relevant (NR)
    NR
    NR
    NR
    Help
    Removal of non-target species [Show more]

    Removal of non-target species

    Benchmark. Removal of features or incidental non-targeted catch (by-catch) through targeted fishery, shellfishery or harvesting at a commercial or recreational scale. Further detail

    Evidence

    Direct, physical impacts are assessed through the abrasion and penetration of the seabed pressures, while this pressure considers the ecological or biological effects of by-catch. Species in this biotope, including the characterizing species Polydora ciliata, may be damaged or directly removed by static or mobile gears that are targeting other species (see abrasion and penetration pressures). Loss of Polydora species and the mat of tubes would alter the character of the biotope resulting in re-classification. Loss of Polydora spp. and the associated epifauna would alter the physical structure of the habitat and result in the loss of the ecosystem functions such as secondary production performed by these species.

    Sensitivity assessment. Removal of the characterizing species would result in the biotope being lost or re-classified. Thus, the biotope is considered to have a resistance of None to this pressure and to have High resilience, resulting in the sensitivity being judged as Medium.

    None
    High
    High
    High
    Help
    High
    High
    Medium
    High
    Help
    Medium
    High
    Medium
    High
    Help

    Bibliography

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    2. Blanchard, M., 2009. Recent expansion of the slipper limpet population (Crepidula fornicata) in the Bay of Mont-Saint-Michel (Western Channel, France). Aquatic Living Resources, 22 (1), 11-19. DOI https://doi.org/10.1051/alr/2009004

    3. Blanchard, M., 1997. Spread of the slipper limpet Crepidula fornicata (L.1758) in Europe. Current state and consequences. Scientia Marina, 61, Supplement 9, 109-118. Available from: http://scimar.icm.csic.es/scimar/index.php/secId/6/IdArt/290/

    4. Bohn, K., Richardson, C. & Jenkins, S., 2012. The invasive gastropod Crepidula fornicata: reproduction and recruitment in the intertidal at its northernmost range in Wales, UK, and implications for its secondary spread. Marine Biology, 159 (9), 2091-2103. DOI https://doi.org/10.1007/s00227-012-1997-3

    5. Bohn, K., Richardson, C.A. & Jenkins, S.R., 2015. The distribution of the invasive non-native gastropod Crepidula fornicata in the Milford Haven Waterway, its northernmost population along the west coast of Britain. Helgoland Marine Research, 69 (4), 313.

    6. Bohn, K., Richardson, C.A. & Jenkins, S.R., 2013b. The importance of larval supply, larval habitat selection and post-settlement mortality in determining intertidal adult abundance of the invasive gastropod Crepidula fornicata. Journal of Experimental Marine Biology and Ecology, 440, 132-140. DOI https://doi.org/10.1016/j.jembe.2012.12.008

    7. Bohn, K., Richardson, C.A. & Jenkins, S.R., 2013a. Larval microhabitat associations of the non-native gastropod Crepidula fornicata and effects on recruitment success in the intertidal zone. Journal of Experimental Marine Biology and Ecology, 448, 289-297. DOI https://doi.org/10.1016/j.jembe.2013.07.020

    8. Borja, A., Franco, J. & Perez, V., 2000. A marine biotic index to establish the ecological quality of soft-bottom benthos within European estuarine and coastal environments. Marine Pollution Bulletin, 40 (12), 1100-1114.

    9. Boulcott, P. & Howell, T.R.W., 2011. The impact of scallop dredging on rocky-reef substrata. Fisheries Research (Amsterdam), 110 (3), 415-420.

    10. Bradshaw, C., Veale, L.O., Hill, A.S. & Brand, A.R., 2002. The role of scallop-dredge disturbance in long-term changes in Irish Sea benthic communities: a re-analysis of an historical dataset. Journal of Sea Research, 47, 161-184. DOI https://doi.org/10.1016/S1385-1101(02)00096-5

    11. 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.

    12. Callier, M. D., McKindsey, C.W. & Desrosiers, G., 2007. Multi-scale spatial variations in benthic sediment geochemistry and macrofaunal communities under a suspended mussel culture. Marine Ecology Progress Series, 348, 103-115.

    13. Como, S. & Magni, P., 2009. Temporal changes of a macrobenthic assemblage in harsh lagoon sediments. Estuarine, Coastal and Shelf Science, 83 (4), 638-646.

    14. Connor, D.W., Allen, J.H., Golding, N., Howell, K.L., Lieberknecht, L.M., Northen, K.O. & Reker, J.B., 2004. The Marine Habitat Classification for Britain and Ireland. Version 04.05. ISBN 1 861 07561 8. In JNCC (2015), The Marine Habitat Classification for Britain and Ireland Version 15.03. [2019-07-24]. Joint Nature Conservation Committee, Peterborough. Available from https://mhc.jncc.gov.uk/

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    17. De Montaudouin, X., Blanchet, H. & Hippert, B., 2018. Relationship between the invasive slipper limpet Crepidula fornicata and benthic megafauna structure and diversity, in Arcachon Bay. Journal of the Marine Biological Association of the United Kingdom, 98 (8), 2017-2028. DOI https://doi.org/10.1017/s0025315417001655

    18. 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.

    19. 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.

    20. 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.

    21. Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.

    22. Gittenberger, A. & Van Loon, W.M.G.M., 2011. Common marine macrozoobenthos species in the Netherlands, their characteristics and sensitivities to environmental pressures. GiMaRIS Report no 2011.08. DOI: https://doi.org/10.13140/RG.2.1.3135.7521

    23. Grassle, J.F. & Grassle, J.P., 1974. Opportunistic life histories and genetic systems in marine benthic polychaetes. Journal of Marine Research, 32, 253-284.

    24. Green, J., 1961. A biology of Crustacea. London: H.F. & G. Witherby Ltd. 180 pp.

    25. 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.

    26. 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.

    27. 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.

    28. 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.

    29. Hansen, B. W., Stenalt, E., Petersen, J.K. & Ellegaard, C., 2002. Invertebrate re-colonisation in Mariager Fjord (Denmark) after severe hypoxia. I. Zooplankton and settlement. Ophelia 56 (3), 197-213.

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    Citation

    This review can be cited as:

    De-Bastos, E.S.R., Hill, J.M., Lloyd, K.A., & Watson, A., 2023. Polydora sp. tubes on moderately exposed sublittoral soft rock. 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. [cited 25-05-2024]. Available from: https://marlin.ac.uk/habitat/detail/247

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    Last Updated: 13/12/2023