Protodorvillea kefersteini and other polychaetes in impoverished circalittoral mixed gravelly sand

Summary

UK and Ireland classification

Description

In coarse gravelly or shelly sand sometimes with a slight mud content, along open coasts in depths of 10 to 30 m, and in shallower offshore areas, an impoverished community characterised by Protodorvillea kefersteini may be found. This biotope has a number of other species associated with it including Nemertea spp., Chaetozone zetlandicaPrionospio cirriferaGlycera lapidumAmpelisca spinipes and numerous other polychaete species all occurring at low abundances. The polychaete Sabellaria spinulosa can also be found in low numbers in this biotope. This biotope has been reported in the North Sea along the Norfolk/Lincolnshire coast located in and around marine aggregate dredging areas. This biotope may be quite variable both spatially and temporally in terms of community structure and also sediment type, which is often borderline between the SCS complex and the SMx complex. (Information from JNCC, 2022). 

Depth range

10-20 m, 20-30 m

Additional information

-

Listed By

Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

The biotope description and characterizing species are taken from JNCC (2015).  This biotope occurs in coarse gravelly or shelly sand sometimes with a slight mud content, and is characterized by an impoverished community defined by Protodorvillea kefersteini with species such as Nemertea spp., Caulleriella zetlandica, Minuspio cirrifera (now Prionospio cirrifera), Glycera lapidum and Ampelisca spinipes all occurring at low abundances. The polychaete Sabellaria spinulosa is also found in low numbers in this biotope but does not form reefs. The sensitivity assessments focus on the sediments which are a key factor structuring the biological assemblage and the polychaetes Protodorvillea kefersteini and Glycera lapidum and the amphipod Ampelisca spinipes.  More information is available for these species than other taxa and the sensitivity of these taxa is considered to represent the sensitivity of the biotope.

Resilience and recovery rates of habitat

The major factor driving the presence of interstitial fauna such as Hesionura elongata is likely to be sediment type (Nybakken, 2001). Sediment type and faunal abundance and diversity are intrinsically linked (Basford et al., 1990; Seiderer & Newell 1999; Cooper et al., 2011), and this is most relevant to interstitial fauna, as these require sediments of a certain grain size that is large enough to enable fauna to inhabit the voids between grains (Nybakken 2001). Food sources are limited for interstitial fauna characterizing this biotope and availability of food is likely to be an important factor influencing recovery. The characterizing species include active predators and deposit feeding detritivores. Predators, such as Hesionura elongata, are known to feed on other interstitial fauna and various infaunal invertebrate species (MES, 2008). 

The characterizing polychaete Protodorvillea is  small (1-3 cm)  and lives in a soft mucous tube under stones, in empty serpulid tubes and in shallow burrows under the surface of muddy sand. It is a carnivore that feeds on small invertebrates at the sediment surface (MES, 2010) and has limited mobility (MES, 2010). The lifespan of this genus is about 1 year and sexual maturity is at about 4-6 months. There is little information on the breeding season or fecundity. After fertilisation, the embryos are brooded before release as planktotrophic larvae and juveniles. The short lifespan, relatively rapid growth rate and larval dispersal phase suggest that this genus has a high recoverability (MES, 2010).

The predatory polychaete, Glycera spp. are longer-lived and monotelic, having a single breeding period towards the end of their life.  Recovery may occur through migration and this species may persist in disturbed sediments through their ability to burrow (Klawe & Dickie, 1952). Glycera spp. have a high potential rate of recolonization of sediments, but the relatively slow growth-rate and long-lifespan suggests that recovery of biomass following initial recolonization by post-larvae is likely to take several years (MES, 2010).  

Sardá et al. (1999) tracked annual cycles within a Spisula community in Bay of Blanes (north west Mediterranean sea, Spain) for 4 years.  Protodorvillea kefersteini exhibited Spring recruitment and the population  persisted throughout the year. Following dredging of subtidal sands in Summer and Autumn to provide material for beach nourishment in the Bay of Blanes, (north west Mediterranean sea, Spain) recovery was tracked by Sardá et al. (2000). Recolonization in the dredged habitats was rapid, for some bivalve and polychaete species but Protodorvillea kefersteini had not recovered within two years (Sardá et al., 2000).

The amphipod genus Ampelisca have some life history traits that allow them to recovery quickly where populations are disturbed. They do not produce large numbers of offspring but reproduce regularly and the larvae are brooded, giving them a higher chance of survival within a suitable habitat than free-living larvae. Ampelisca has a short lifespan and reaches sexual maturity in a matter of months allowing a population to recover abundance and biomass in a very short period of time (MES, 2008). Experimental studies have shown Ampelisca abdita  to be an early colonizer, in large abundances of defaunated sediments where local populations exist to support recovery (McCall, 1977) and Ampelisca abdita  have been shown to migrate to, or from, areas to avoid unfavourable conditions (Nichols & Thompson, 1985). Ampelisca sp. are very intolerant of oil contamination and the recovery of the Ampelisca populations in the fine sand community in the Bay of Morlaix took up to 15 years following the Amoco Cadiz oil spill, probably due to the amphipods' low fecundity, lack of pelagic larvae and the absence of local unperturbed source populations (Poggiale & Dauvin, 2001). 

Where impacts also alter the sedimentary habitat, recovery of the biotope will also depend on recovery of the habitat to the former condition to support the characteristic biological assemblage. Recovery of sediments will be site-specific and will be influenced by currents, wave action and sediment availability (Desprez, 2000). Except in areas of mobile sands, the process tends to be slow (Kenny & Rees, 1996; Desprez, 2000 and references therein).  Boyd et al., (2005) found that in a site where sands and gravels were subject to long-term extraction (25 years), extraction scars were still visible after six years and sediment characteristics were still altered in comparison with reference areas, with ongoing effects on the biota.

Resilience assessment. Where resistance is ‘None’ or ‘Low’ and an element of habitat recovery is required, resilience is assessed as ‘Medium’ (2-10 years), based on evidence from aggregate recovery studies in similar habitats including Boyd et al. (2005); Where resistance of the characterizing species is ‘Low’ or ‘Medium’ and the habitat has not been altered, resilience is assessed as ‘High’ as it is likely that the biotope would be considered representative and hence recovered after two years although some parameters such as species richness, abundance and biotopes may be altered . Recovery of the seabed from severe physical disturbances that alter sediment character may also take up to 10 years or longer (Le Bot et al., 2010), although extraction of gravel may result in more permanent changes and this will delay recovery. 

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

Limited evidence was found on the effect of changes in temperature and resistance is inferred from the characterizing species ranges. 

Hesionura elongata occurs in the Canary Islands and Caribbean, which suggests a resistance of higher water temperatures than around UK and Irish seas (Brito et al., 2005; Miloslavich et al., 2010). Protodorvillea kefersteini is found in the North Atlantic to the  North Sea and English Channel, Mediterranean and Black Sea (Marine Species Identification Portal).

Bamber & Spencer (1984) observed that Cauleriella zetlandica were dominant species in an area affected by thermal discharge in the River Medway estuary. Cauleriella zetlandica were dominant resident fauna present in high abundances. The species is clearly tolerant of temperature fluctuations as the sediments were exposed to the passage of a temperature front of approximately 10 oC between heated effluent and estuarine waters during the tidal cycles.

Sensitivity assessment. This assessment relies on limited evidence and utilises global species distribution records to assess sensitivity (with the exception of Caulleriella zetlandica)  and so confidence is low. As all characterizing species occur in water temperatures greater than they are likely to experience in the UK, biotope resistance and resilience are assessed as ‘High’ and the biotope is considered to be ‘Not Sensitive’. There is low confidence associated with this assessment as limited evidence was available.

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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

Limited evidence was returned on the effect of changes in temperature and resistance is inferred from the species range. Hesionura elongata has been identified in samples from water ranging from 7.3-24°C (OBIS, 2016). Protodorvillea kefersteini is found in the North Atlantic to the North Sea and English Channel, Mediterranean and Black Sea (Marine Species Identification Portal).

Sensitivity assessment. Limited evidence was available and this assessment is based on non-peer reviewed literature on species range. A 5°C decrease in temperature for one month period is likely to impact the characterizing species in winter months and therefore resistance is ‘Medium’, resilience is ‘High’ and Sensitivity is ‘Low’. 

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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

The biotope occurs in ‘full’ salinity conditions. An increase in one MNCR salinity category to hypersaline conditions is likely to cause mortality of characterizing species. Resistance is ‘None’, Resilience is ‘High (following restoration of habitat conditions)’ and sensitivity is assessed as ‘High’. This assessment is assessed based on distribution and confidence is 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

This biotope occurs in full salinity habitats. A change at the pressure benchmark is assessed as a decrease from full to reduced salinity(18-30 ppt)

Degraer et al. (2006) report that Hesionura elongata was found in greatest abundance outside the near coastal zone (in samples from across the Belgium part of the North Sea). This suggests that the species is likely to occur in greater abundance in habitats with full salinity compared to variable salinity or reduced. Moulaert et al. (2008) also found that species communities in which Hesionura elongata was an indicator species were only present >16 km from the coast and displayed a positive correlation with increasing salinity.

Sensitivity assessment. Resistance is assessed as ‘Medium’ as Hesionura elongata  and other characterizing species may decrease in abundance. Resilience is assessed as ‘High’ and sensitivity is assessed as ‘Low’.

Medium
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Low
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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

No information on tidal streams was presented in the biotope description from JNCC (2015). This biotope occurs in gravelly or shelly sands. Sands are less cohesive than mud sediments and a change in water flow at the pressure benchmark may alter sediment transport patterns within the biotope.  Hjulström (1939), concluded that fine sand (particle diameter of 0.3-0.6 mm) was easiest to erode and required a mean velocity of 0.2 m/s. Erosion and deposition of particles greater than 0.5 mm require a velocity > 0.2 m/s to alter the habitat. The topography of this habitat is shaped by currents and wave action that influence the formation of ripples in the sediment. Specific fauna may be associated with troughs and crests of these bedforms which may form following an increase in water flow, or disappear following a reduction in flow.

The characterizing species are infaunal and may be tolerant of changes in water flow. Glycera spp. are found in areas with strong tidal streams where sediments are mobile (Roche et al. 2007) and in extremely sheltered areas (Connor et al. 2004). 

Sensitivity assessment. This biotope probably occurs in areas subject to moderately strong water flows that are a key factor maintaining the clean sand habitat. Changes in water flow may alter the topography of the habitat and may cause some shifts in abundance. However, a change at the pressure benchmark (increase or decrease)  is unlikely to affect biotopes that occur in mid-range flows and biotope sensitivity is therefore assessed as ‘High’ and resilience is assessed as ‘High’ so that the biotope is considered to be ‘Not sensitive’.

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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

This biotope does not occur in the intertidal, and consequently an increase in emergence is considered not relevant to this biotope.

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

As this biotope occurs in circalittoral habitats it is not directly exposed to the action of breaking waves.  Associated polychaete species that burrow are protected within the sediment. They and other associated species may be indirectly affected by changes in water movement where these impact the supply of food or larvae or other processes. No specific evidence was found to assess this pressure. 

Sensitivity assessment. The abundance of characterizing species is likely to be unaffected or increase in areas where fine sediment is removed and coarse sediment is present. However, abundance is likely to decrease in areas where fine sediment is deposited. Under the pressure benchmark levels which consider <5% change, Resistance is assessed as  ‘High’ and resilience as ‘High’ and the biotope is assessed as ‘Not Sensitive’.

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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.

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.

Contamination at levels greater than the pressure benchmark may adversely influence the biotope. Suchanek (1993) reviewed the effects of oil spills on marine invertebrates and concluded that, in general, on soft sediment habitats, infaunal polychaetes, bivalves and amphipods were particularly affected. The 1969 West Falmouth Spill of Grade 2 diesel fuel, documented by Sanders (1978), illustrates the effects of hydrocarbons in a sheltered habitat with a soft mud/sand substrata (Suchanek, 1993). The entire benthic fauna was eradicated immediately following the spill and remobilization of oil that continued for a period >1 year after the spill contributed to much greater impact upon the habitat than that caused by the initial spill. Effects are likely to be prolonged as hydrocarbons incorporated within the sediment by bioturbation will remain for a long time, owing to slow degradation under anoxic conditions. Oil covering the surface and within the sediment would prevent oxygen transport to the infauna and promote anoxia as the infauna utilise oxygen during respiration. Although this study investigates impacts on an estuarine biotope the impact on benthic infauna communities is likely to be similar in shallow sandbank biotopes.

The amphipods, Ampelisca sp. are very intolerant of oil contamination and the recovery of the Ampelisca populations in the fine sand community in the Bay of Morlaix took up to 15 years following the Amoco Cadiz oil spill (Poggiale & Dauvin, 2001). Other species present in the bivalve are more tolerant f hydrocarbons. Levell et al., (1989) classified Cauleriella sp. and Glycera sp, were all tolerant of elevated hydrocarbons and increased in abundance in the transitional zones along pollution gradients.

 

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.

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

Insufficient information was available in relation to characterizing species to assess this pressure. Limited evidence is available on other infauna species. Beasley & Fowler (1976) and Germain et al., (1984) examined the accumulation and transfers of radionuclides in Hediste diversicolor from sediments contaminated with americium and plutonium derived from nuclear weapons testing and the release of liquid effluent from a nuclear processing plant. Both concluded that the uptake of radionuclides by Hediste diversicolor was small. Beasley & Fowler (1976) found that Hediste diversicolor accumulated only 0.05% of the concentration of radionuclides found in the sediment. Both also considered that the predominant contamination pathway for Hediste diversicolor was from the interstitial water.

Sensitivity assessment: There is insufficient information available on the biological effects of radionuclides to comment further upon the intolerance of characterizing species to radionuclide contamination. Assessment is given as ‘No Evidence’.

No evidence (NEv)
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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.

Some, all be it limited evidence was returned by searches on activated carbon (AC). AC is utilised in some instances to effectively remove organic substances from aquatic and sediment matrices. Lillicrap et al. (2015) demonstrate that AC may have physical effects on benthic dwelling organisms at environmentally relevant concentrations at remediated sites.

 

Not Assessed (NA)
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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

Limited evidence was returned on effects of decreased dissolved oxygen concentrations on the characterizing species.

All meiofauna have some sensitivity to extended hypoxia, although more mobile nematode species are able to emigrate into the water column in high numbers where they survive (Wetzel et al., 2013). Emigration is likely to increase predation risk. Although evidence on characterizing species is lacking, densities of meiofauna populations are likely to be lower under prolonged anoxia (Moodley et al., 1997).  

Riedel et al. (2012) assessed the response of benthic macrofauna to hypoxia advancing to anoxia in the Mediterranean. The hypoxic and anoxic conditions were created for 3-4 days in a box that enclosed in-situ sediments. In general, molluscs were more resistant than polychaetes, with 90% surviving hypoxia and anoxia, whereas only 10% of polychaetes survived, the exposed Glycera spp. died. Sensitivity may vary between species and populations, or due to factors such as temperature as Glycera alba was reported to be able to tolerate periods of anoxia resulting from inputs of organic material from a wood pulp and paper mill in Loch Eil (Scotland) (Blackstock & Barnes, 1982). 

Sensitivity assessment. Due to the limited evidence confidence in this assessment is low. A reduction in meiofauna populations is likely if deoxygenation persisted for long periods, but this is unlikely due to high water flow. As some species are likely to emigrate or maintain aerobic metabolism under low dissolved oxygen conditions, Resistance is assessed as ‘Medium’, Resilience is ‘High’ and Sensitivity is assessed as ‘Low’.

Medium
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Low
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Nutrient enrichment [Show more]

Nutrient enrichment

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

Evidence

Meiofauna respond to nutrient enrichment. The distribution of different meiofauna assemblages has been identified as a good tool for detecting short-term responses of the benthic domain to nutrient enrichment from sources such as river discharge (Semprucci et al., 2015). In the Bay of Cadiz, Spain, abundance of meiofauna was seven times higher in the presence of macroalgae (Bohorquez et al., 2013).

Sensitivity assessment. As the benchmark levels comply with WFD criteria for good status, Resistance is ‘High’, Resilience is ‘High’ and the biotope is 'Not sensitive' at the benchmark level.

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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

Protodorvillea kefersteini was identified as a ‘progressive’ species, i.e. one that shows increased abundance under slight organic enrichment (Leppakoski, 1975, cited in Gray, 1979; Hiscock et al., 2004). Protodorvillea kefersteini can become very plentiful in organically enriched habitats (Warwick et al., 1986), this species was very abundant in the vicinity of a sewage outfall at Kircaldy (S.C. Hull pers. comm).  Protodorvillea  kefersteini were dominant species in muddy,  organically enriched sediments (organic content approximately 25%) located about 100 and 500 m from fish farm cages, in a bay in Corsica, France (Terlizzi et al., 2010). Similarly, this species was also dominant in sediments close to fish farms in Greek bays where organic matter and nitrogen content had increased.

Glycera lapidum has been categorized through expert judgement and literature review as AMBI Group III- Species tolerant to excess organic matter enrichment. This species may occur under normal conditions, but populations are stimulated by organic enrichment (slight unbalance situations) (Borja et al., 2000, validated by Gittenberger & van Loon, 2011).

Sensitivity assessment.  Protodorvillea kefersteini is tolerant of organic enrichmentAt the pressure benchmark organic inputs are considered likely to represent a food subsidy and are unlikely to significantly affect the structure of the biological assemblage or impact the physical habitat. Biotope sensitivity is therefore assessed as ‘High’ and resilience as ‘High’ (by default) and the biotope is therefore considered to be ‘Not sensitive’.

 

 

No evidence (NEv)
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No evidence (NEv)
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No evidence (NEv)
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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
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Very Low
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High
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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

This biotope is only found in circalittoral sand habitats, the characterizing polychaete species burrow or live interstitially within the sediment and would not be able to survive if the substratum type was changed to either a soft rock or hard artificial type. Consequently, the biotope would be lost altogether if such a change occurred. 

Sensitivity assessment.  The Resistance to this change is ‘None’, and the Resilience is assessed as ‘Very low’, due to the long-term nature of a change in substratum.  The biotope is assessed to have a ‘High’ Sensitivity to this pressure at the benchmark. 

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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

Increase in gravel content within the Folk classes is unlikely to negatively impact the characterizing species Hesionura elongata. Increases in finer sand or silt are likely to reduce abundance of Hesionura elongata  as this species is found in greater abundance in sediments with larger grain sizes, and decreases in abundance in fine sediments. Moulaert & Hostens (2007) found that higher gravel content and sediment grain size was a key environmental factor determining the presence of Hesionura elongata. Desprez (2000) found that a change of habitat to fine sands from coarse sands and gravels (from deposition of screened sand following aggregate extraction) changed the biological communities present.

Sensitivity assessment.  Sediment changes are likely to alter the composition of the biological assemblage leading to biotope reclassification. Biotope resistance is therefore assessed as ‘Low’ (as some species may remain), biotope resilience is assessed as ‘Very low' (the pressure is a permanent change) and biotope sensitivity is assessed as ‘High’.

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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

A number of studies assess the impacts of aggregate extraction on sand and gravel habitats. Extraction would remove the infauna that may be present in this biotope. Recovery of sediments will be site-specific and will be influenced by currents, wave action and sediment availability (Desprez, 2000). Except in areas of mobile sands, the process tends to be slow (Kenny & Rees, 1996; Desprez, 2000 and references therein).  Boyd et al., (2005) found that in a site subject to long-term extraction (25 years), extraction scars were still visible after six years and sediment characteristics were still altered in comparison with reference areas with ongoing effects on the biota. The strongest currents are unable to transport gravel. A further implication of the formation of these depressions is a local drop in current strength associated with the increased water depth, resulting in deposition of finer sediments than those of the surrounding substrate (Desprez et al., 2000 and references therein). See the physical change pressure for assessment

Sensitivity assessment. Resistance is assessed as ‘None’ as extraction of the sediment swill remove the characterizing and associated species present. Resilience is assessed as ‘Medium’ as some species may require longer than two years to re-establish (see resilience section) and sediments may need to recover (where exposed layers are different). Biotope sensitivity is therefore assessed as ‘Medium’.

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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

No evidence. The characterizing species are infaunal and likely to be protected from abrasion, although movement of sediments may damage a proportion of the population. Ampelisca spp. inhabit surface tubes; surface abrasion will damage these and expose the amphipods to increased predation. Kaiser and Spencer (1994) found that fish preyed more heavily on Ampelisca spinipes following beam trawling as damage to the tubes exposed the amphipods. As Ampelisca is not a key characterizing species,  biotope resistance based on polychaetes is assessed as 'Medium' and resilience as 'High', so that biotope sensitivity is considered to be 'Low'.

Medium
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Low
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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

This biotope may be an impoverished version of a biotope such as SS.SCS.CCS.MedLumVen or similar, which has been subject to storm disturbance or other physical disturbance such as aggregate dredging (JNCC, 2015). this suggests that the species present are either resistant of disturbance or are early colonizers following disturbance.

Protodorvillea kefersteini is soft-bodied and therefore vulnerable to damage by physical abrasion. However, its environmental position as burrowing interstitial species should provide a high degree of protection from activities that lead to surface abrasion only. Similarly, as a small polychaete species, living  infaunally  and capable of burrowing rapidly, Hesionura elongata is also  likely to withstand physical disturbance caused by bottom towed fishing gears (such as otter or beam trawls) (Vanosmael et al., 1982; Bolam et al., 2014).  Experiments in shallow, wave disturbed areas, using a toothed, clam dredge, found that some polychaete taxa without external protection and with a carnivorous feeding mode were enhanced by fishing. Protodorvillea kefersteini was one of these: large increases in abundance in samples were detected post dredging and persisting over 90 days. The passage of the dredge across the sediment floor will have killed or injured some organisms that will then be exposed to potential predators/scavengers (Frid et al., 2000; Veale et al., 2000) providing a food source to mobile scavengers including these species. 

 In a coarse gravelly substratum exposed to high current velocities the crab Cancer pagurus was observed to dig pits, approximately 30 cm in diameter and 10 cm deep. Experiments were conducted to identify macrobenthic recolonization processes and differences in abundance between pits and unmanipulated areas. Protodorvillia kefersteini (McIntosh) (Polychaeta) showed a rapid increase in abundance at 21 days after disturbance (Thrush, 1986).

Ampelisca brevicornis has been categorised through expert and literature review as AMBI fisheries Group I-Species very sensitive to fisheries in which the bottom is disturbed. Their populations do not easily recover (Gittenberger & van Loon, 2011). This assessment is probably equally applicable to Ampelisca spinipesGlycera alba and Glycera lapidum were categorised as AMBI fisheries Group III- second-order opportunistic species, which are sensitive to fisheries in which the bottom is disturbed. Their populations recover relatively quickly however and benefit from the disturbance, causing their population sizes to increase significantly in areas with intense fisheries (Gittenberger & van Loon, 2011). Glycera lapidum is present in the biotope SS.SCS.ICS.Glap which is an impoverished biotope type subject to sediment destabilisation by wave action (Connor et al., 2004).

Capasso et al. (2010), compared benthic survey datasets from 1895 and 2007 for an area in the English Channel. Although methodological differences limit direct comparison, the datasets appear to show that large, fragile urchin species including Echinus esculentus, Spatangus purpureus and Psammechinus miliaris and larger bivalves had decreased in abundance. Small, mobile species such as amphipods and small errant and predatory polychaetes (Nephtys, Glycera, Lumbrineris) appeared to have increased (Capasso et al. 2010).  The area is subject to beam trawling and scallop dredging and the observed species changes would correspond with predicted changes following physical disturbance (Capasoo et al., 2010). 

Sensitivity assessment. Evidence is limited but the biological assemblage present in this biotope is characterized by species that are likely to be relatively tolerant of penetration and disturbance of the sediments, with the exception of the amphipod Ampelisca spinipes. Either species are robust or buried within sediments or are adapted to habitats with frequent disturbance (natural or anthropogenic) and recover quickly. Biotope resistance is assessed as ‘Medium’ based on the characterizing polychaetes as some species will be displaced and may be predated or injured and killed. Biotope resilience is assessed as ‘High’ as most species will recover rapidly. Biotope sensitivity is therefore assessed as ‘Low’.

Medium
High
High
High
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High
High
High
High
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Low
High
High
High
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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

No direct evidence was found to assess this pressure, the characterizing polychaetes live infaunally and are predatory and may not be directly impacted by either a decrease or increase in suspended solids.

Amphipods are tolerant of high turbidity and gather suspended sediment for the construction of tubes. Mills (1967) reported that feeding by Ampelisca vadorum and Ampelisca abdita was initiated by the turbidity of the water surrounding the tubes. However, the feeding structures of suspension feeders such as Ampelisca sp. may become clogged by large increases in suspended sediment or feeding may be terminated, compromising growth. A decrease in turbidity and hence increased light penetration may result in increased phytoplankton production and hence indirectly enhance food supply for suspension feeders, including Ampelisca spp (although phytoplankton are not a major part of the diet). Therefore, reduced turbidity may be beneficial. In areas of high suspended sediment, a decrease may result in improved habitat quality and recruitment. However, a decrease in suspended organic particles in some areas may reduce food availability resulting in lower growth or reduced energy for reproduction and less silt available for tube production. 

Sensitivity assessmentBased on the characterizing polychaetes biotope resistance is assessed as 'High' and resilience as 'High' so that the biotope is assessed as 'Not sensitive'.

 

High
Low
NR
NR
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High
High
High
High
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Not sensitive
Low
Low
Low
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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

No evidence.

No evidence (NEv)
NR
NR
NR
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No evidence (NEv)
NR
NR
NR
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No evidence (NEv)
NR
NR
NR
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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

No evidence.

No evidence (NEv)
NR
NR
NR
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No evidence (NEv)
NR
NR
NR
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No evidence (NEv)
NR
NR
NR
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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

No evidence was returned on the impact of litter on characterizing species for this biotope, although studies show impacts from ingestion of micro plastics by sub surface deposit feeding worms (Arenicola marina) and toxicants present in cigarette butts have been shown to impact the burrowing times and cause DNA damage in ragworms Hediste diversicolor.

Litter, in the form of cigarette butts has been shown to have an impact on Ragworms. Hediste diversicolor showed increased burrowing times, 30% weight loss and a  >2 fold increase in DNA damage when exposed to water with toxicants (present in cigarette butts) in quantities 60 fold lower than reported from urban run-off (Wright et al., 2015). Studies are limited on impacts of litter on infauna and this UK study suggests health of infauna populations are negatively impacted by this pressure.

Studies of sediment dwelling, sub surface deposit feeding worms, a trait shared by species abundant in this biotope, showed negative impacts from ingestion of micro plastics. For instance, Arenicola marina ingests micro plastics that are present within the sediment it feeds within. Wright et al. (2013) carried out a lab study that displayed presence of micro plastics (5% UPVC) significantly reduced feeding activity when compared to concentrations of 1% UPVC and controls. As a result, Arenicola marina showed significantly decreased energy reserves (by 50%), took longer to digest food, and decreased bioturbation levels. These effects would be likely to impact colonisation of sediment by other species, reducing diversity in the biotopes the species occurs within. Wright et al. (2013) also present a case study based on their results, that in the intertidal regions of the Wadden Sea, where Arenicola marina is an important ecosystem engineer, Arenicola marina could ingest 33mᵌ; of micro plastics a year.

Sensitivity assessment. ‘No evidence’ was returned to complete a sensitivity assessment, however, both microplastics and the toxicants present in cigarette butts are likely to have negative impacts on the characterizing species.

Not Assessed (NA)
NR
NR
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Not assessed (NA)
NR
NR
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Not assessed (NA)
NR
NR
NR
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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 was found on effects of electric and magnetic fields on the characterizing species.

Electric and magnetic fields generated by sources such as marine renewable energy device/array cables may alter behaviour of predators and affect infauna populations. Evidence is limited and occurs for electric and magnetic fields below the benchmark levels, confidence in evidence of these effects is very low.

Field measurements of electric fields at North Hoyle wind farm, North Wales recorded 110µ V/m (Gill et al., 2009). Modelled results of magnetic fields from typical subsea electrical cables, such as those used in the renewable energy industry produced magnetic fields of between 7.85 and 20 µT (Gill et al., 2009; Normandeau et al., 2012). Electric and magnetic fields smaller than those recorded by in field measurements or modelled results were shown to create increased movement in thornback ray Raja clavata and attraction to the source in catshark Scyliorhinus canicular (Gill et al., 2009).

Flatfish species which are predators of many polychaete species including dab Limanda limanda and sole Solea solea have been shown to decrease in abundance in a wind farm array or remain at distance from wind farm towers (Vandendriessche et al., 2015; Winter et al., 2010). However, larger plaice increased in abundance (Vandendriessche et al., 2015). There have been no direct causal links identified to explain these results.

Sensitivity assessment.No evidence’ was available to complete a sensitivity assessment, however, responses by flatfish and elasmobranchs suggest changes in predator behaviour are possible. There is currently no evidence but effects may occur on predator prey dynamics as further marine renewable energy devices are deployed, these are likely to be over small spatial scales and not impact the biotope.

No evidence (NEv)
NR
NR
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No evidence (NEv)
NR
NR
NR
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No evidence (NEv)
NR
NR
NR
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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

Species within the biotope can probably detect vibrations caused by noise and in response may retreat in to the sediment for protection. However, at the benchmark level the community is unlikely to be sensitive to noise and therefore is ‘Not sensitive’.

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

Introduction of light or shading

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

Evidence

All characterizing species live in the sediment and do not rely on light levels directly to feed or find prey so limited direct impact is expected.  Most species will respond to the shading caused by the approach of a predator, however, their visual acuity is probably very low. Even then, additional disturbance, such as an electronic flash, caused the retraction of palps and cirri and cessation of all activity for some minutes. Visual disturbance, in the form of direct illumination during the species' active period at night, may therefore result in loss of feeding opportunities, which may compromise growth and reproduction.

As this biotope is not characterized by the presence of primary producers it is not considered that shading would alter the character of the habitat directly. More general changes to the productivity of the biotope may, however, occur. Beneath shading structures there may be changes in microphytobenthos abundance, which would affect food resources (Tait & Dipper, 1998).

Shading will prevent photosynthesis leading to death or migration of sediment microalgae altering sediment cohesion and food supply to higher trophic levels. The impact of these indirect effects is difficult to quantify.

Sensitivity assessment. Based on the direct impact, biotope Resistance is assessed as ‘High’ and Resilience is assessed as ‘High’ (by default). The biotope Sensitivity is considered to be ‘Not sensitive’.

High
Low
NR
NR
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High
High
High
High
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Not sensitive
Low
Low
Low
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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

This biotope is reported in offshore waters (JNCC, 2015) and this pressure is considered 'Not relevant'.

Not relevant (NR)
NR
NR
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Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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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 interaction with bottom towed fishing gears and moorings are addressed under ‘surface abrasion’.

Not relevant (NR)
NR
NR
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Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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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

Characterizing species may have some, limited, visual perception. As they live in the sediment the species will most probably not be impacted at the pressure benchmark.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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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

Characterizing species are not cultivated or translocated. This pressure is 'Not relevant'.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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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

The American slipper limpet Crepidula fornicata was introduced to the UK and Europe in the 1870s from the Atlantic coasts of North America with imports of the eastern oyster Crassostrea virginica. It was recorded in Liverpool in 1870 and the Essex coast in 1887-1890. It has spread through expansion and introductions along the full extent of the English Channel and into the European mainland (Blanchard, 1997, 2009; Bohn et al., 2012, 2013a, 2013b, 2015; De Montaudouin et al., 2018; Hinz et al., 2011; McNeill et al., 2010; Powell-Jennings & Calloway, 2018; Preston et al., 2020; Stiger-Pouvreau & Thouzeau, 2015).

Crepidula fornicata is recorded from shallow, sheltered bays, lagoons and estuaries or the sheltered sides of islands, in variable salinity (18 to 40) although it prefers ca 30 (Tillin et al., 2020). 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; Powell-Jennings & Calloway, 2018; Preston et al., 2020; Stiger-Pouvreau & Thouzeau, 2015; Tillin et al., 2020).  

High densities of Crepidula fornicata cause ecological impacts on sedimentary habitats. The species can smother the seabed in shallow bays, changing and modifying the habitat structure. At high densities, the species physically smothers the sediment, and the resultant build-up of silt, pseudofaeces, and faeces is deposited and trapped within the bed (Tillin et al., 2020, Fitzgerald, 2007, Blanchard, 2009, Stiger-Pouvreau & Thouzeau, 2015). The biodeposition rates of Crepidula are extremely high and once deposited, form an anoxic mud, making the environment suitable for other species, including most infauna (Stiger-Pouvreau & Thouzeau, 2015, Blanchard, 2009). For example, in fine sands, the community is replaced by a reef of slipper limpets, that provide hard substrata for sessile suspension-feeders (e.g., sea squirts, tube worms and fixed shellfish), while mobile carnivorous microfauna occupy species between or within shells, resulting in a homogeneous Crepidula dominated habitat (Blanchard, 2009). Blanchard (2009) suggested the transition occurred and became irreversible at 50% cover of the limpet. De Montaudouin et al. (2018) suggested that homogenization occurred above a threshold of 20-50 Crepidula /m2.

Impacts on the structure of benthic communities will depend on the type of habitat that Crepidula colonizes. De Montaudouin & Sauriau (1999) reported that in muddy sediment dominated by deposit-feeders, species richness, abundance and biomass increased in the presence of high densities of Crepidula (ca 562 to 4772 ind./m2), in the Bay of Marennes-Oléron, presumably because the Crepidula bed provided hard substrata in an otherwise sedimentary habitat. In medium sands, Crepidula density was moderate (330-1300 ind./ m2) but there was no significant difference between communities in the presence of Crepidula.  Intertidal coarse sediment was less suitable for Crepidula with only moderate or low abundances (11 ind./m2) and its presence did not affect the abundance or diversity of macrofauna. However, there was a higher abundance of suspension–feeders and mobile Crustacea in the absence of Crepidula (De Montaudouin & Sauriau, 1999). The presence of Crepidula as an ecosystem engineer has created a range of new niche habitats, reducing biodiversity as it modifies habitats (Fitzgerald, 2007). De Montaudouin et al. (1999) concluded that Crepidula did not influence macroinvertebrate diversity or density significantly under experimental conditions, on fine sands in Arcachon Bay, France. De Montaudouin et al. (2018) noted that the limpet reef increased the species diversity in the bed, but homogenised diversity compared to areas where the limpets were absent. In the Milford Haven Waterway (MHW), the highest densities of Crepidula were found in areas of sediment with hard substrata, e.g., mixed fine sediment with shell or gravel or both but, while Crepidula density increased as gravel cover increased in the subtidal, the reverse was found in the intertidal (Bohn et al., 2015). Bohn et al. (2015) suggested that high densities of Crepidula in high-energy environments were possible in the subtidal but not the intertidal.  Hinz et al. (2011) reported a substantial increase in the occurrence of Crepidula off the Isle of Wight, between 1958 and 2006, at a depth of ca 60 m, on hard substrata (gravel, cobbles, and boulders), swept by strong tidal streams. Presumably, Crepidula is more tolerant of tidal flow than the oscillatory flow caused by wave action (Tillin et al., 2020). 

Sensitivity assessment. The above evidence suggests that Crepidula fornicata could colonize coarse sediment habitats in the subtidal, typical of this biotope. Bohn et al. (2015) demonstrated that Crepidula had a preference for gravelly habitats, while De Montaudouin & Sauriau (1999) and Bohn et al. (2015) noted that Crepidula densities were low in intertidal coarse sediments. Therefore, if Crepidula colonized this biotope, it would probably modify the habitat and its associated community due to the introduction of Crepidula shell biomass, silt, pseudofaeces and faeces (Blanchard, 2009; Tillin et al., 2020), as occurs in maerl gravels (Grall & Hall-Spencer, 2003) resulting in the loss of the biotope. This is a disturbed, impoverished habitat, in which storms may mobilise the sediment (JNCC, 2022), which may mitigate or prevent colonization by Crepidula at high densities, although Crepidula has been recorded from areas of strong tidal streams (Hinz et al., 2011). However, Crepidula reduced the density of suspension feeders and mobile Crustacea in coarse sediment even at low densities (De Montaudouin & Sauriau, 1999).  Therefore, resistance is assessed as 'Low'. Resilience is assessed as 'Very low' as it would require the removal of Crepidula, probably by artificial means. Hence, sensitivity is assessed as 'High'. Crepidula has not yet been reported to occur in this biotope so the confidence in the assessment is 'Low' and further evidence is required. 

Low
Low
NR
NR
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Very Low
High
High
High
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High
Low
NR
NR
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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

No evidence was found.

No evidence (NEv)
NR
NR
NR
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Not relevant (NR)
NR
NR
NR
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No evidence (NEv)
NR
NR
NR
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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

No characterizing species are targeted directly by fishing activities at a commercial or recreational scale, this pressure is therefore ‘Not relevant’.

Not relevant (NR)
NR
NR
NR
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Not relevant (NR)
NR
NR
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Not relevant (NR)
NR
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NR
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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

Species within the biotope are not functionally dependent on each other, although biological interactions will play a role in structuring the biological assemblage through predation and competition. Removal of species would also reduce the ecological services provided by these species such as secondary production and nutrient cycling.

Sensitivity assessment. Species within the biotope are relatively sedentary or slow moving although the infaunal position may protect some burrowing species from removal. Biotope resistance is, therefore, assessed as ‘Low’ and resilience as ‘High’ as the habitat is likely to be directly affected by removal and some species will recolonize rapidly.  Therefore, sensitivity is assessed as 'Low'. Some variability in species recruitment, abundance and composition is natural and, therefore, a return to a recognisable biotope should occur within 2 years. Repeated chronic removal would, however, impact recovery.

Low
Low
NR
NR
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High
High
Low
Medium
Help
Low
Low
Low
Low
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Bibliography

  1. Alexander, D., Colcombe, A., Chambers, C. & Herbert, R.J.H. 2014. Conceptual Ecological Modelling of Shallow Sublittoral Coarse Sediment Habitats to Inform Indicator Selection. Marine Ecological Surveys Ltd - A report for the Joint Nature Conservation Committee, JNCC Report No: 520

  2. Bamber, R.N. & Spencer, J.F. 1984. The benthos of a coastal power station thermal discharge canal. Journal of the Marine Biological Association of the United Kingdom, 64, 603-623.

  3. Beasley, T.M. & Fowler, S.W., 1976. Plutonium and Americium: uptake from contaminated sediments by the polychaete Nereis diversicolor. Marine Biology, 38, 95-100.

  4. Bijkerk, R., 1988. Ontsnappen of begraven blijven: de effecten op bodemdieren van een verhoogde sedimentatie als gevolg van baggerwerkzaamheden: literatuuronderzoek: RDD, Aquatic ecosystems.

  5. Blackstock, J. & Barnes, M., 1982. The Loch Eil project: biochemical composition of the polychaete, Glycera alba (Müller), from Loch Eil. Journal of Experimental Marine Biology and Ecology, 57 (1), 85-92.

  6. 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

  7. 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/

  8. 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

  9. 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.

  10. 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

  11. 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

  12. Bohorquez, J., Papaspyrou, S., Yufera, M., van Bergeijk, S.A., Garcia-Robledo, E., Jimenez-Arias, J.L., Bright, M. & Corzo, A., 2013. Effects of green macroalgal blooms on the meiofauna community structure in the Bay of Cadiz. Marine Pollution Bulletin, 70 (1-2), 10-17.

  13. Bolam, S.G., Coggan, R.C., Eggleton, J., Diesing, M. & Stephens, D., 2014. Sensitivity of macrobenthic secondary production to trawling in the English sector of the Greater North Sea: A biological trait approach. Journal of Sea Research, 85, 162-177.

  14. 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.

  15. Boyd, S., Limpenny, D., Rees, H. & Cooper, K., 2005. The effects of marine sand and gravel extraction on the macrobenthos at a commercial dredging site (results 6 years post-dredging). ICES Journal of Marine Science: Journal du Conseil, 62 (2), 145-162.

  16. Brito, M.C., Martin, D. & Nunez, J., 2005. Polychaetes associated to a Cymodocea nodosa meadow in the Canary Islands: assemblage structure, temporal variability and vertical distribution compared to other Mediterranean seagrass meadows. Marine Biology, 146 (3), 467-481.

  17. Capasso, E., Jenkins, S., Frost, M. & Hinz, H., 2010. Investigation of benthic community change over a century-wide scale in the western English Channel. Journal of the Marine Biological Association of the United Kingdom, 90 (06), 1161-1172.

  18. 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/

  19. Cooper, K., Ware, S., Vanstaen, K. & Barry, J., 2011. Gravel seeding - A suitable technique for restoring the seabed following marine aggregate dredging? Estuarine, Coastal and Shelf Science, 91 (1), 121-132.

  20. De Montaudouin, X. & Sauriau, P.G., 1999. The proliferating Gastropoda Crepidula fornicata may stimulate macrozoobenthic diversity. Journal of the Marine Biological Association of the United Kingdom, 79, 1069-1077. DOI https://doi.org/10.1017/S0025315499001319

  21. De Montaudouin, X., Andemard, C. & Labourg, P-J., 1999. Does the slipper limpet (Crepidula fornicata L.) impair oyster growth and zoobenthos diversity ? A revisited hypothesis. Journal of Experimental Marine Biology and Ecology, 235, 105-124.

  22. 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

  23. Degraer, S., Wittoeck, J., Appeltans, W., Cooreman, K., Deprez, T., Hillewaert, H., Hostens, K., Mees, J., Vanden Berghe, E. & Vincx, M., 2006. The macrobenthos atlas of the Belgian part of the North Sea. Belgian Science Policy, Brussels.

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Citation

This review can be cited as:

Tillin, H.M. & Watson, A., 2023. Protodorvillea kefersteini and other polychaetes in impoverished circalittoral mixed gravelly sand. In Tyler-Walters H. Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 09-12-2024]. Available from: https://www.marlin.ac.uk/habitat/detail/1115

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Last Updated: 30/08/2023