The Marine Life Information Network

The honeycomb worm, Sabellaria spinulosa has the greatest geographical range of all the sabellariids, according to current records, encompassing Iceland, the Skagerrak, and the Kattegat, the North Sea, the English Channel, the North East Atlantic, the Mediterranean, the Wadden Sea, and the Indian Ocean, (Achari, 1974; Riesen & Reise, 1982; Reise & Schubert,1987; Hayward & Ryland, 1998; Foster-Smith, 2000; Collins, 2005). It does not form reefs across most of its geographic range (Jackson & Hiscock, 2008), although Sabellaria spinulosa reefs are known from all North Atlantic European coasts, except the Baltic and the waters of the Kattegat and Skagerrak (OSPAR, 2010). In the Mediterranean Sea, the most common reef-forming sabellid worm is Sabellaria alveolata, although extensive Sabellaria spinulosa reefs have been found in the Adriatic Sea (Gravina et al., 2018).

Sensitivity assessment. Under the middle and high emission and extreme scenarios seawater temperatures are expected to temperatures rise by 3-5°C to potential southern summer temperatures of 22 - 24°C and northern summer temperatures of 19 - 21°C. No empirical evidence was found for the temperature tolerance of Sabellaria spinulosa; nevertheless, biogeographic distribution is often a good predictor of temperature tolerance (Jeffree & Jeffree, 1994), and its widespread distribution suggests that it is tolerant to temperature variations (Gibb et al., 2014).  Extensive Sabellaria spinulosa reefs are found off the coast of Vieste, Italy, in the Adriatic Sea (Gravina et al., 2018), where water temperatures range from 12 – 28°C (www.seatemperature.org), which suggests that this biotope will be able to withstand the predicted temperature changes for the end of this century. Therefore, this biotope is assessed as having a ‘High’ resistance to ocean warming.  Resilience is assessed as ‘High, as no recovery is necessary. This biotope is assessed as ‘Not sensitive’ under the middle and high emission and extreme scenarios

The honeycomb worm, Sabellaria spinulosa has the greatest geographical range of all the sabellariids, according to current records, encompassing Iceland, the Skagerrak, and the Kattegat, the North Sea, the English Channel, the North East Atlantic, the Mediterranean, the Wadden Sea, and the Indian Ocean, (Achari, 1974; Riesen & Reise, 1982; Reise & Schubert,1987; Hayward & Ryland, 1998; Foster-Smith, 2000; Collins, 2005). It does not form reefs across most of its geographic range (Jackson & Hiscock, 2008), although Sabellaria spinulosa reefs are known from all North Atlantic European coasts, except the Baltic and the waters of the Kattegat and Skagerrak (OSPAR, 2010). In the Mediterranean Sea, the most common reef-forming sabellid worm is Sabellaria alveolata, although extensive Sabellaria spinulosa reefs have been found in the Adriatic Sea (Gravina et al., 2018).

Sensitivity assessment. Under the middle and high emission and extreme scenarios seawater temperatures are expected to temperatures rise by 3-5°C to potential southern summer temperatures of 22 - 24°C and northern summer temperatures of 19 - 21°C. No empirical evidence was found for the temperature tolerance of Sabellaria spinulosa; nevertheless, biogeographic distribution is often a good predictor of temperature tolerance (Jeffree & Jeffree, 1994), and its widespread distribution suggests that it is tolerant to temperature variations (Gibb et al., 2014).  Extensive Sabellaria spinulosa reefs are found off the coast of Vieste, Italy, in the Adriatic Sea (Gravina et al., 2018), where water temperatures range from 12 – 28°C (www.seatemperature.org), which suggests that this biotope will be able to withstand the predicted temperature changes for the end of this century. Therefore, this biotope is assessed as having a ‘High’ resistance to ocean warming.  Resilience is assessed as ‘High, as no recovery is necessary. This biotope is assessed as ‘Not sensitive’ under the middle and high emission and extreme scenarios

The honeycomb worm, Sabellaria spinulosa has the greatest geographical range of all the sabellariids, according to current records, encompassing Iceland, the Skagerrak, and the Kattegat, the North Sea, the English Channel, the North East Atlantic, the Mediterranean, the Wadden Sea, and the Indian Ocean, (Achari, 1974; Riesen & Reise, 1982; Reise & Schubert,1987; Hayward & Ryland, 1998; Foster-Smith, 2000; Collins, 2005). It does not form reefs across most of its geographic range (Jackson & Hiscock, 2008), although Sabellaria spinulosa reefs are known from all North Atlantic European coasts, except the Baltic and the waters of the Kattegat and Skagerrak (OSPAR, 2010). In the Mediterranean Sea, the most common reef-forming sabellid worm is Sabellaria alveolata, although extensive Sabellaria spinulosa reefs have been found in the Adriatic Sea (Gravina et al., 2018).

Sensitivity assessment. Under the middle and high emission and extreme scenarios seawater temperatures are expected to temperatures rise by 3-5°C to potential southern summer temperatures of 22 - 24°C and northern summer temperatures of 19 - 21°C. No empirical evidence was found for the temperature tolerance of Sabellaria spinulosa; nevertheless, biogeographic distribution is often a good predictor of temperature tolerance (Jeffree & Jeffree, 1994), and its widespread distribution suggests that it is tolerant to temperature variations (Gibb et al., 2014).  Extensive Sabellaria spinulosa reefs are found off the coast of Vieste, Italy, in the Adriatic Sea (Gravina et al., 2018), where water temperatures range from 12 – 28°C (www.seatemperature.org), which suggests that this biotope will be able to withstand the predicted temperature changes for the end of this century. Therefore, this biotope is assessed as having a ‘High’ resistance to ocean warming.  Resilience is assessed as ‘High, as no recovery is necessary. This biotope is assessed as ‘Not sensitive’ under the middle and high emission and extreme scenarios

Marine heatwaves due to increased air-sea heat flux are predicted to occur more frequently, last for longer and at increased intensity by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018). Whilst there are no laboratory studies on the upper thermal limit of Sabellaria spinulosa, its biogeographical distribution suggests that, similar to the reef-forming Sabellaria alveolata, this species may be tolerant to a wide range of temperatures.

Muir et al. (2016) found that Sabellaria alveolata was able to adapt to a step-change increase in temperature from 15°C to 25°C, however, over the long term (60 days), changes in lipid composition possibly suggested a stress response. Whilst high temperatures may cause stress, they rarely appear to lead to mortality, as when summer temperatures were increased from 18°C to 23°C in Scottish populations of Sabellaria alveolata, mortality remained the same as controls, except when coupled with high levels of chlorine (Last et al., 2016). Similarly, when the density of tube occupancy was used as a proxy for mortality by Muir et al. (2016), an increase in temperatures from 15°C to 25°C in populations from Scotland to North Africa did not lead to a decrease in occupancy in four out of the five populations tested, with a decrease in occupancy only observed for the Bay of Biscay population, which the authors attributed to the poor state of the reef.

Sensitivity Assessment. Under the middle emission scenario, if heatwaves occurred every three years, with a maximum intensity of 2°C for 80 days by the end of this century, this could lead to summer sea temperatures reaching up to 24°C in southern England. Under the high emission scenario, if heatwaves occur every two years by the end of this century, reaching a maximum intensity of 3.5°C for 120 days, this could lead to the heatwave lasting the entire summer with temperatures reaching up to 26.5°C. Sabellaria spinulosa is known to form extensive reefs in the Adriatic Sea (Gravina et al., 2018), where water temperatures reach up to 28°C (www.seatemperature.org) (see Global warming). Whilst empirical evidence of how Sabellaria spinulosa would react to exposure to a marine heatwave is lacking, it is likely that this species would be able to tolerate marine heatwaves at both these levels. Therefore, this species is likely to be able to cope with future marine heatwaves expected for the end of this century, and under both the middle and high emission scenarios, and resistance has been assessed as ‘High’. As no recovery is likely necessary, recovery has been assessed as ‘High’, leading to an assessment of ‘Not sensitive’ for this biotope. 

Marine heatwaves due to increased air-sea heat flux are predicted to occur more frequently, last for longer and at increased intensity by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018). Whilst there are no laboratory studies on the upper thermal limit of Sabellaria spinulosa, its biogeographical distribution suggests that, similar to the reef-forming Sabellaria alveolata, this species may be tolerant to a wide range of temperatures.

Muir et al. (2016) found that Sabellaria alveolata was able to adapt to a step-change increase in temperature from 15°C to 25°C, however, over the long term (60 days), changes in lipid composition possibly suggested a stress response. Whilst high temperatures may cause stress, they rarely appear to lead to mortality, as when summer temperatures were increased from 18°C to 23°C in Scottish populations of Sabellaria alveolata, mortality remained the same as controls, except when coupled with high levels of chlorine (Last et al., 2016). Similarly, when the density of tube occupancy was used as a proxy for mortality by Muir et al. (2016), an increase in temperatures from 15°C to 25°C in populations from Scotland to North Africa did not lead to a decrease in occupancy in four out of the five populations tested, with a decrease in occupancy only observed for the Bay of Biscay population, which the authors attributed to the poor state of the reef.

Sensitivity Assessment. Under the middle emission scenario, if heatwaves occurred every three years, with a maximum intensity of 2°C for 80 days by the end of this century, this could lead to summer sea temperatures reaching up to 24°C in southern England. Under the high emission scenario, if heatwaves occur every two years by the end of this century, reaching a maximum intensity of 3.5°C for 120 days, this could lead to the heatwave lasting the entire summer with temperatures reaching up to 26.5°C. Sabellaria spinulosa is known to form extensive reefs in the Adriatic Sea (Gravina et al., 2018), where water temperatures reach up to 28°C (www.seatemperature.org) (see Global warming). Whilst empirical evidence of how Sabellaria spinulosa would react to exposure to a marine heatwave is lacking, it is likely that this species would be able to tolerate marine heatwaves at both these levels. Therefore, this species is likely to be able to cope with future marine heatwaves expected for the end of this century, and under both the middle and high emission scenarios, and resistance has been assessed as ‘High’. As no recovery is likely necessary, recovery has been assessed as ‘High’, leading to an assessment of ‘Not sensitive’ for this biotope. 

Increasing levels of CO₂ in the atmosphere have led to the average pH of sea surface waters dropping from 8.25 in the 1700s to 8.14 in the 1990s (Jacobson, 2005). There is no direct evidence of the impact of ocean acidification on Sabellaria spinulosa or any species from the family Sabellariidae. Unlike the tube-dwelling, calcifying polychaetes from the family Serpulidae, Sabellaria spinulosa does not form its tubes through calcification. Tubes are formed from coarse sand and shell grains cemented together with an adhesive protein cement secreted by the worm (Lisco et al., 2017).

Non-calcifying polychaetes thought to be less sensitive than many other taxa to ocean acidification.  When non-calcifying polychaetes were transplanted from control to low pH areas, they showed evidence of either adaptation or acclimation to their conditions (Calosi et al., 2013). There is some evidence that sperm may be affected by ocean acidification at levels expected in the high emission scenario, with percentage sperm motility (Schlegel et al., 2014) and sperm velocity (Campbell et al., 2014) decreasing in the polychaetes Galeolaria caespitosa and Arenicola marina, leading to a decrease in sperm fertility success (Campbell et al., 2014). Reduced sperm fertility and hence recruitment, may lead to some population-level effects. However, at natural CO₂ vents, the abundance of polychaetes either remained the same (Kroeker et al., 2011) or increased (Garrard et al., 2014, Vizzini et al., 2017). Most species of polychaetes generally exhibit high fecundity and are free spawning (Ramirez-Llodra, 2002), which may help them maintain population levels, even with a decrease in fertilization success. Fecundity in Sabellaria spinulosa expected to be high and is thought to be similar to that of Sabellaria alveolata (Jackson & Hiscock, 2008), which releases an average of 100,000 eggs into the water column during a spawning cycle (Dubois, 2003).

Sensitivity Assessment. Direct evidence of the impact of ocean acidification on Sabellaria spinulosa is lacking. However, in general, non-calcifying polychaetes appear to be tolerant. Therefore, it is likely that the characterizing species of this biotope will show a ‘High’ resistance to a decrease in pH, even though ocean acidification has been shown to lead to negative impacts on polychaete fertilization success under experimental conditions (Campbell et al., 2014, Schlegel et al., 2014). Hence, based on the evidence available, under both the middle and high emission scenarios the biotope is assessed as ‘High’ resistance to ocean acidification, and ‘High’ resilience, leading to an assessment of ‘Not sensitive’ at the benchmark level.

Increasing levels of CO₂ in the atmosphere have led to the average pH of sea surface waters dropping from 8.25 in the 1700s to 8.14 in the 1990s (Jacobson, 2005). There is no direct evidence of the impact of ocean acidification on Sabellaria spinulosa or any species from the family Sabellariidae. Unlike the tube-dwelling, calcifying polychaetes from the family Serpulidae, Sabellaria spinulosa does not form its tubes through calcification. Tubes are formed from coarse sand and shell grains cemented together with an adhesive protein cement secreted by the worm (Lisco et al., 2017).

Non-calcifying polychaetes thought to be less sensitive than many other taxa to ocean acidification.  When non-calcifying polychaetes were transplanted from control to low pH areas, they showed evidence of either adaptation or acclimation to their conditions (Calosi et al., 2013). There is some evidence that sperm may be affected by ocean acidification at levels expected in the high emission scenario, with percentage sperm motility (Schlegel et al., 2014) and sperm velocity (Campbell et al., 2014) decreasing in the polychaetes Galeolaria caespitosa and Arenicola marina, leading to a decrease in sperm fertility success (Campbell et al., 2014). Reduced sperm fertility and hence recruitment, may lead to some population-level effects. However, at natural CO₂ vents, the abundance of polychaetes either remained the same (Kroeker et al., 2011) or increased (Garrard et al., 2014, Vizzini et al., 2017). Most species of polychaetes generally exhibit high fecundity and are free spawning (Ramirez-Llodra, 2002), which may help them maintain population levels, even with a decrease in fertilization success. Fecundity in Sabellaria spinulosa expected to be high and is thought to be similar to that of Sabellaria alveolata (Jackson & Hiscock, 2008), which releases an average of 100,000 eggs into the water column during a spawning cycle (Dubois, 2003).

Sensitivity Assessment. Direct evidence of the impact of ocean acidification on Sabellaria spinulosa is lacking. However, in general, non-calcifying polychaetes appear to be tolerant. Therefore, it is likely that the characterizing species of this biotope will show a ‘High’ resistance to a decrease in pH, even though ocean acidification has been shown to lead to negative impacts on polychaete fertilization success under experimental conditions (Campbell et al., 2014, Schlegel et al., 2014). Hence, based on the evidence available, under both the middle and high emission scenarios the biotope is assessed as ‘High’ resistance to ocean acidification, and ‘High’ resilience, leading to an assessment of ‘Not sensitive’ at the benchmark level.

Sea level rise is occurring through a combination of thermal expansion and ice melt.  Sea levels have risen 1-3 mm/yr. in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). This biotope generally occurs between 10–30 m depth in the UK, although Sabellaria spinulosa reefs have been recorded from 10–50 m depth in Europe (OSPAR, 2010), therefore an increase in depth of between 50–107 cm is unlikely to have large implications for this biotope.

Sabellaria spinulosa reefs occur on stable circalittoral mixed sediment in moderately exposed or sheltered environments with strong tidal currents. Sabellaria spinulosa reefs generally occur in areas of high sediment disturbance (OSPAR, 2010).  Understanding how sea-level rise will affect exposure or tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site-dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude. The effects of sea-level rise and increased wave action may be increased further due to storms and storm surges.  IPCC (2019) note that the frequency of extreme sea-level events (e.g. due to storms) are predicted to increase as sea-level rises, however, there is no consensus on the future storm and, hence, wave climate around UK coasts (Mossman et al., 2015, Lowe et al., 2018, Palmer et al., 2018).

Sensitivity assessment. This habitat occurs from 10-30 m although Sabellaria spinulosa reefs can be found at depths of up to 50 m. Any change to the habitat in terms of its exposure or tidal currents cannot be evaluated at the current time, although evidence suggests that changes to tidal currents and tidal amplitude with sea-level rise will be site-specific. Therefore, under the available evidence, resistance to sea-level rise has been assessed as ‘High’ for both the middle (50 cm) and high (70 cm) emission scenario, and the extreme scenario (107 cm). As no recovery is deemed necessary, resilience has been assessed as ‘High’, and therefore this biotope has been classified as ‘Not sensitive’ to sea-level rise at each of the benchmarks albeit with ‘Low’ confidence.

Sea level rise is occurring through a combination of thermal expansion and ice melt.  Sea levels have risen 1-3 mm/yr. in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). This biotope generally occurs between 10–30 m depth in the UK, although Sabellaria spinulosa reefs have been recorded from 10–50 m depth in Europe (OSPAR, 2010), therefore an increase in depth of between 50–107 cm is unlikely to have large implications for this biotope.

Sabellaria spinulosa reefs occur on stable circalittoral mixed sediment in moderately exposed or sheltered environments with strong tidal currents. Sabellaria spinulosa reefs generally occur in areas of high sediment disturbance (OSPAR, 2010).  Understanding how sea-level rise will affect exposure or tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site-dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude. The effects of sea-level rise and increased wave action may be increased further due to storms and storm surges.  IPCC (2019) note that the frequency of extreme sea-level events (e.g. due to storms) are predicted to increase as sea-level rises, however, there is no consensus on the future storm and, hence, wave climate around UK coasts (Mossman et al., 2015, Lowe et al., 2018, Palmer et al., 2018).

Sensitivity assessment. This habitat occurs from 10-30 m although Sabellaria spinulosa reefs can be found at depths of up to 50 m. Any change to the habitat in terms of its exposure or tidal currents cannot be evaluated at the current time, although evidence suggests that changes to tidal currents and tidal amplitude with sea-level rise will be site-specific. Therefore, under the available evidence, resistance to sea-level rise has been assessed as ‘High’ for both the middle (50 cm) and high (70 cm) emission scenario, and the extreme scenario (107 cm). As no recovery is deemed necessary, resilience has been assessed as ‘High’, and therefore this biotope has been classified as ‘Not sensitive’ to sea-level rise at each of the benchmarks albeit with ‘Low’ confidence.

Sea level rise is occurring through a combination of thermal expansion and ice melt.  Sea levels have risen 1-3 mm/yr. in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). This biotope generally occurs between 10–30 m depth in the UK, although Sabellaria spinulosa reefs have been recorded from 10–50 m depth in Europe (OSPAR, 2010), therefore an increase in depth of between 50–107 cm is unlikely to have large implications for this biotope.

Sabellaria spinulosa reefs occur on stable circalittoral mixed sediment in moderately exposed or sheltered environments with strong tidal currents. Sabellaria spinulosa reefs generally occur in areas of high sediment disturbance (OSPAR, 2010).  Understanding how sea-level rise will affect exposure or tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site-dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude. The effects of sea-level rise and increased wave action may be increased further due to storms and storm surges.  IPCC (2019) note that the frequency of extreme sea-level events (e.g. due to storms) are predicted to increase as sea-level rises, however, there is no consensus on the future storm and, hence, wave climate around UK coasts (Mossman et al., 2015, Lowe et al., 2018, Palmer et al., 2018).

Sensitivity assessment. This habitat occurs from 10-30 m although Sabellaria spinulosa reefs can be found at depths of up to 50 m. Any change to the habitat in terms of its exposure or tidal currents cannot be evaluated at the current time, although evidence suggests that changes to tidal currents and tidal amplitude with sea-level rise will be site-specific. Therefore, under the available evidence, resistance to sea-level rise has been assessed as ‘High’ for both the middle (50 cm) and high (70 cm) emission scenario, and the extreme scenario (107 cm). As no recovery is deemed necessary, resilience has been assessed as ‘High’, and therefore this biotope has been classified as ‘Not sensitive’ to sea-level rise at each of the benchmarks albeit with ‘Low’ confidence.

No empirical evidence was found for the temperature tolerance of Sabellaria spinulosa; nevertheless, its widespread distribution suggests that it is tolerant to temperature variations (Gibb et al., 2014). Sabellaria spinulosa has the greatest geographical range of all the sabellariids, according to current records, encompassing Iceland, the Skagerrak and the Kattegat, the North Sea, the English Channel, the northeast Atlantic, the Mediterranean, the Wadden Sea and the Indian Ocean, (Achari, 1974; Riesen & Reise, 1982; Reise & Schubert,1987; Hayward & Ryland, 1998; Foster-Smith, 2000; Collins, 2005).

Sabellaria spinulosa occurs north to the Arctic and is therefore considered tolerant of a decrease in temperature at the pressure benchmark. This conclusion is supported by observations made on oyster grounds in the River Crouch throughout the severe winter of 1962–1963 that Sabellaria spinulosa appeared unaffected by the cold. The mean daily temperature was recorded at a depth of 1 fathom (1.8m) below low water (equinoctial spring tide) and the lowest temperature recorded was -1.8°C (Crisp, 1964). At Penmon in Bangor, Sabellaria spinulosa also appeared not to suffer from the low temperatures and live individuals were readily found (Crisp, 1964). 

Sensitivity assessment. Given the widespread distribution of Sabellaria spinulosa, it is unlikely that this species is sensitive to temperature variations at the pressure benchmark. Resistance is therefore assessed as ‘High’ and resilience is assessed as ‘High’ (no impact to recover from) so that all the Sabellaria biotopes within this group are assessed as ‘Not Sensitive’.

No evidence for the range of physiological tolerances to salinity changes was found for Sabellaria spinulosa by Gibb et al., (2014).  As reefs are largely subtidal they are less exposed to hypersaline conditions resulting from coastal brine discharge and natural evaporation (lagoons).  There is, therefore, no direct or indirect evidence for sensitivity to an increase in salinity and this element of the pressure is not assessed. 

Sensitivity assessment. No evidence was found for tolerance of hypersaline conditions and sensitivity to this benchmark is not assessed based on lack of evidence.

No evidence for the range of physiological tolerances to decreases in salinity was found for Sabellaria spinulosa by Gibb et al. (2014).  The sensitivity assessment made in that report was therefore based on recorded habitat preferences, as described below. 

Sabellaria spinulosa does not seem to occur in very low salinity areas (Holt et al., 1998) but has been recorded from estuaries including the Crouch, Mersey (Killeen & Light, 2000) and the Thames (Limpenny et al., 2010).  Buhs & Reise (1997) surveyed 12 channel systems in the Wadden Sea and found that Sabellaria spinulosa reefs occurred in the northern tidal inlets which experienced salinity levels ranging from 28 to 30 psu.  There is some speculation (Foster-Smith & Hendrick, 2003) that Mcintosh (1922) misidentified samples of Sabellaria spinulosa as the congener Sabellaria alveolata from the Humber estuarine population (Holt et al., 1998).  These records indicate that reduced and variable salinities can be tolerated to some extent but the paucity of records suggests that areas of reduced salinity do not provide optimal habitat.

Sensitivity assessment. As the salinity tolerances of Sabellaria spinulosa are unclear the potential impact of salinity change, at the pressure benchmark, is uncertain.  The reported distribution of Sabellaria spinulosa from fully marine to estuarine habitats does suggest some tolerance of changes in salinity although a decrease in salinity at the extreme of the pressure benchmark (reduction to variable salinity 18-35 ppt or reduced salinity 18-30 ppt) may result in extensive losses.  Resistance is therefore assessed as ‘Low’ (loss of 25-75% of extent).  Reef resilience (following habitat recovery) is considered to be “Medium’ (2-10 years).  Sensitivity, based on combined resistance and recovery, is therefore assessed as ‘Medium’.

Sabellaria spinulosa tend to occur in areas of high water movement where larvae, tube building materials and food particles are suspended and transported (Jones et al., 2000).  The relative importance of tidal versus wave induced movements to support reefs is, however, unclear (Holt et al., 1998). There is currently limited in-situ data on the specific water flow tolerances of Sabellaria spinulosa, although colonies have been found in areas with sedimentary bed forms that suggest current velocities in the range of 0.5 m/s to 1.0 m/s (Mistakidis, 1956; Jones et al., 2000; Davies et al., 2009).  In the southern North Sea close to the coast of England, Sabellaria spinulosa reefs have been recorded in areas exposed to peak spring tidal flows of 1.0 m/s (Pearce et al., 2014). Davies et al. (2009) also found through laboratory experiments with Sabellaria spinulosa in tanks that increased the water flow to an average of 0.03 m/s is adequate to begin distribution of the sediment rain from the airlift throughout the tank and that doubling the water flow to almost 0.07 m/s further improved particle distribution throughout the tank.  It is therefore likely that Sabellaria spinulosa will exist in habitats with a water flow anywhere above 0.07 m/s so that particles are suspended and distributed for the use of tube building and feeding. 

Tillin (2010) used logistic regression to develop statistical models that indicate how the probability of occurrence of the congener Sabellaria alveolata changes over environmental gradients within the Severn Estuary.  The model predicted response surfaces were derived for each biotope for each of the selected habitat variables, using logistic regression.  From these response surfaces the optimum habitat range for each biotope could be defined based on the range of each environmental variable where the probability of occurrence, divided by the maximum probability of occurrence, is 0.75 or higher.  These results identify the range for each significant variable where the habitat is most likely to occur.  The modelled ranges should be interpreted with caution and apply to the Severn Estuary alone (which experiences large tidal ranges, high currents and extremely high suspended sediment loads and is therefore distinct from many other estuarine systems).  However, these ranges do provide some useful information on environmental tolerances.  The models indicate that for subtidal Sabellaria alveolata the maximum optimal current speed (the range in which it is most likely to occur) ranges from 1.26-2.46 m/s and the optimal mean current speed ranges from 0.5-1.22 m/s.  Although not directly applicable to Sabellaria spinulosa this data suggests that tube-building Sabellariids are able to occur within a broad range of current speeds.

In cases of reduced water flow, Sabellaria spinulosa is likely to suffer a reduction in the supply of suspended food and particles that are integral for growth and repair.  A long-term decrease in water flow may reduce the viability of populations by limiting growth and tube building. No evidence was found for threshold levels relating to impact.

Sensitivity assessment. The range of flow tolerances recorded (0.5 m/s to 1 m/s cited by Jones et al., 2000; Braithwaite et al., 2006; Davies et al., 2009) suggest that the worms have a broad tolerance of different flow levels.  Tillin (2010) modelled optimal flow speeds of 0.5-1.22 m/s for the congener Sabellaria alveolata.  The worms may retract into tubes to withstand periods of high flows at spring tides and some non-lethal reduction in feeding efficiency and growth rate may occur at the edge of the range.  Similarly, a reduction in flow may reduce the supply of tube-building materials and food but again, given the range of reported tolerances a change at the pressure benchmark is not considered to result in mortality.  Resistance is therefore assessed as ‘High’ and resilience as ‘High’ (no impact to recover from).  All the biotopes within this biotope group are therefore considered to be ‘Not sensitive’.

Changes in emergence are not relevant to this biotope, which is restricted to subtidal habitats.

No empirical evidence was found to assess this pressure.  Intertidal Sabellaria spinulosa are directly exposed to breaking waves and water movements generated by waves can also potentially affect subtidal Sabellaria spinulosa reefs.  At depth, the motion from surface waves becomes oscillatory and produces back-and-forth water movement at the seabed (Dubois et al., 2006).  In sublittoral habitats, water movements are likely to provide sand and food particles that are necessary for Sabellaria spinulosa to build tubes, feed and subsequently grow and develop.  

Sensitivity assessment. As Sabellaria spinulosa reefs are robust, stable structures that are present subtidally in naturally disturbed environments and areas with high water flow, changes (decrease or increase) in wave height at the pressure benchmark are not considered to affect reefs.  All biotopes within this group are therefore considered to have ‘High’ resistance to this pressure, resilience is assessed as ‘High’ (no impact to recover from) and all subtidal reef biotopes are considered to be ‘Not Sensitive’.  Intertidal populations of Sabellaria spinulosa would be more exposed to the impacts of wave exposure but the corresponding biotopes for these habitats are not included in this assessment.  

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

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

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

No evidence.

This pressure is Not assessed.

Cole et al. (1999) suggest possible adverse effects on marine species below 4 mg/l and probable adverse effects below 2 mg/l. No information was found regarding Sabellaria spinulosa tolerance to changes in oxygenation and this pressure is not assessed due to lack of evidence.

No direct evidence was found to assess this pressure. As the reefs are circalittoral increased nutrient enrichment (eutrophication) would not stimulate the overgrowth of macroalgae on reefs as light penetration is too limited (especially in turbid areas) to allow growth. Enhanced phytoplankton production may increase food supply and increased siltation and deoxygenation from algal blooms is likely to be mitigated by water movements in the areas most suitable for Sabellaria spinulosa reefs. 

Sensitivity assessment. At the pressure benchmark, which refers to the maintenance of 'good' status according to the Water Framework Directive, Sabellaria spinulosa reefs are considered 'Not sensitive'. 

Limited direct evidence was found to assess the effects of this pressure. The presence and enhanced growth of Sabellaria spinulosa adjacent to a sludge dumping area in Dublin (Walker & Rees 1980)  suggest that Sabellaria spinulosa reef biotopes are resistant to a high level of organic enrichment.  Information on the levels of organic matter in Dublin Bay was not provided and so it is unclear how the levels experienced relate to the pressure benchmark. 

Sabellaria spinulosa reefs are found in areas of high water movement (up to 1 m/s from (Pearce et al., 2014, see 'change in water flow' for further details) that would naturally disperse some organic matter preventing accumulation and siltation.  In larger, dense colonies of Sabellaria spinulosa, sand, detritus, and finer faecal materials collect in between worm tubes.  These detritus layers do not interrupt the normal growth of the individuals or of the colony as a whole (Schafer, 1972).  Taking into consideration these points it seems likely that Sabellaria spinulosa are resistant to the deposition of a fine layer of organic materials.

Indirect effects arising from inputs of organic matter are possible where habitat quality and species interactions are altered.  In the Wadden Sea, large subtidal areas of Sabellaria spinulosa reefs have been completely lost since the 1920s.  This decline has been partly attributed to an increase in coastal eutrophication that has favoured blue mussel beds (Dörjes, 1992; Hayward & Ryland, 1998; Benson et al., 2013).  However, a direct causal link has not been established and it is possible that the decline of Sabellaria spinulosa reefs was due to physical damage from fishing activities rather than competitive interactions (Jones et al., 2000).

Sensitivity assessment. Little evidence was found to support this sensitivity assessment.  Habitat preferences for areas of high water movement suggest that organic matter would not accumulate on reefs, limiting exposure to this pressure.  Sabellaria spinulosa and the associated species assemblage (which typically includes attached filter feeders from a number of phyla) is likely to be able to consume extra organic matter.  This conclusion is supported by the enhanced growth rates that have been recorded in the vicinity of sewage disposal areas (Walker & Rees, 1980).  Resistance is therefore assessed as ‘High’ to this pressure and recovery is assessed as ‘High’ (no impact to recover from) and the biotope is considered to be ‘Not Sensitive’ at the pressure benchmark. 

All marine habitats and benthic species are considered to have ‘No Resistance’ to this pressure and to be unable to recover from a permanent loss of habitat.  Sensitivity within the direct spatial footprint of this pressure is, therefore ‘High’.  Although no specific evidence is described confidence in the resistance assessment is ‘High’, due to the incontrovertible nature of this pressure.  Adjacent habitats and species populations may be indirectly affected where meta-population dynamics and trophic networks are disrupted and where the flow of resources e.g. sediments, prey items, loss of nursery habitat etc. is altered. No recovery is predicted to occur.

The introduction of artificial hard substratum or the exposure of natural hard substratum is considered at the pressure benchmark level and it is noted that Sabellaria spinulosa can colonise bedrock and artificial structures (Mistakidis, 1956).  An increase in the availability of hard substratum may, therefore, be beneficial in areas where sedimentary habitats were previously unsuitable for colonisation.  

Sensitivity assessment. Based on reported habitat preferences the species (rather than the biotope) is considered to be ‘Not Sensitive’ as the resulting habitat is suitable for the development of reefs (however these would be classified as a different biotope type).  The resistance of the biotope is, therefore, assessed as None (loss of >75% of extent), resilience is Very low (the pressure is a permanent change) and sensitivity is assessed as High. The more precautionary assessment for the biotope, rather than the species, is presented in the table as it is considered that any change from a sedimentary habitat to a rock reef habitat would alter the biotope classification and hence the more sensitive assessment is appropriate.

Dredging and dumping of sediment and infrastructure developments can lead to changes in sediment character.  The impact of the change will depend on the sediment changes that result.  Foster-Smith (2001b) reported that the best Sabellaria spinulosa reefs identified in an area of the Wash were associated with ground clearly scarred by dredging activities.  It is believed that this was most likely due to a reduction in the overburden of sand resulting in a substratum more suitable for reef formation (Foster-Smith, 2001b).

Sabellaria spinulosa biotopes that occur on mixed sediments are not considered to be affected by a change in sediment type of one Folk class that leads to a change to ‘coarse sediments’ characterized as gravel, sandy gravel or gravelly sand (based on the Long (2006) simplified Folk classification) or a change to sands and muddy sand as this species is found on sands (George & Warwick, 1985).  It should be note, that changes in substratum type and water flows, that resulted in a loss of supply of suitable tube-building materials, would negatively impact this biotope. However, this biotope is considered to be negatively impacted by a change to the fine sediment e.g. a change in the sediment classification to ‘mud and sandy mud’.  This assessment is based on the lack of records of reefs occurring on these sediment types and is likely due to the mobility of the sediment, the lack of sand for tube-building and possibly the re-suspension of fine sediments clogging feeding structures and gills, however, this is assumed rather than based on direct evidence.

Sensitivity assessment.  Based on reported habitat preferences and evidence from Foster-Smith (2001b), where a change in one Folk class results in increased coarseness (e.g. a change to a coarse sediment of gravel, sandy gravel or gravelly sand) then the biotope is considered to be ‘Not Sensitive’ as the resulting habitat is suitable for this species.  However, an increase in fine sediments to the degree that sediments are re-classified as mud or sandy mud would severely reduce habitat suitability.  Therefore, resistance has been assessed as ‘None’, resilience as Very low (the pressure is a permanent change), and sensitivity as High. 

The removal of sediment or substratum down to 30 cm depth is likely to remove the whole Sabellaria spinulosa reef within the extraction footprint.  For example, no reefs were present in the active dredge zone when dredging was taking place at the Hastings Shingle Bank  (Pearce et al., 2007).  Therefore resistance to this pressure is assessed as ‘None’.  However, if suitable substrata were to remain (as specified in the pressure benchmark), reef recovery would depend on larval settlement and survival. 

Hill et al., (2011) reviewed the recoverability of seabed sediments following marine aggregate extraction.  Rapid recovery was reported in areas with high levels of sediment mobility (8 months) which is likely to include the habitat that Sabellaria spinulosa is commonly found in (Holt et al., 1998).  For example, in areas such as the Bristol Channel (where Sabellaria spinulosa is currently distributed), physical traces of dredging that had been carried out in mobile sandy habitats disappeared within a few tidal cycles (Newell et al., 1998).  Similarly, dredge tracks at an area of the North Sea exposed to high levels of wave action disappeared in less than a year (Hill et al., 2011).  With regards to Sabellaria spinulosa specifically, rapid recovery after the cessation of dredging has been observed at high dredging intensity zones in the Hastings Shine Bank area (new establishment observed in less than a year) (Pearce et al., 2007).  

Sensitivity assessment. As Sabellaria spinulosa reefs are present on the surface they will be directly removed by extraction of the sediment, resistance to this pressure is therefore assessed as ‘None’.  Resilience informed by (Pearce et al., 2007) is considered to be ‘Medium’ to allow for the establishment of reef structure and the potential for variable recruitment and this biotope is therefore considered to have ‘Medium’ sensitivity to this pressure.

Sabellaria spinulosa reef biotopes are directly exposed to physical damage that affects the surface.  Gibb et al. (2014) found no direct evidence for impacts of the surface only for Sabellaria spinulosa.  Studies of intertidal reefs of the congener Sabellaria alveolata (Cunningham et al.,1984) have found that the reef recovered within 23 days from the effects of trampling, (i.e. treading, walking or stamping on the reef structures) by repairing minor damage to the worm tube porches.  Severe, localised damage, caused by kicking and jumping on the reef structure, resulted in large cracks between the tubes, and removal of sections (ca 15 x15 x10 cm) of the structure (Cunningham et al., 1984).  Subsequent wave action enlarged the holes or cracks.  However, after 23 days, at one site, one side of the hole had begun to repair, and tubes had begun to extend into the eroded area. 

To address concerns regarding damage from fishing activities in the Wadden Sea, Vorberg (2000) used video cameras to study the effect of shrimp fisheries on Sabellaria alveolata reefs.  The imagery showed that the 3m beam trawl easily ran over a reef that rose to 30 to 40 cm, although the beam was occasionally caught and misshaped on the higher sections of the reef.  At low tide, there were no signs of the reef being destroyed and, although the trawl had left impressions, all traces had disappeared four to five days later due to the rapid rebuilding of tubes by the worms.  The daily growth rate of the worms during the restoration phase was significantly higher than undisturbed growth (undisturbed: 0.7 mm, after removal of 2 cm of surface: 4.4 mm) and indicates that as long as the reef is not completely destroyed recovery can occur rapidly. 

Sabellaria spinulosa reefs are suggested to be more fragile than Sabellaria alveolata (B. Pearce, pers comm, cited from Gibb et al., 2014) and therefore surface abrasion may lead to greater damage and lower recovery rates than observed for Sabellaria alveolata.  Sabellaria spinulosa reefs are often only approximately 10cm thick, surface abrasion can, therefore, severely damage and/or remove a reef (see also evidence from penetration and disturbance of the sunbstratum, below).  No direct observations of reef recovery, through repair, from abrasion were found for Sabellaria spinulosa.

Sensitivity assessment. Based on the evidence discussed above, abrasion at the surface of Sabellaria spinulosa reefs is considered likely to damage the tubes and result in sub-lethal and lethal damage to the worms.  Resistance is therefore assessed as ‘Low’ (loss of 25-75% of tubes and worms within the impact footprint).  Resilience is therefore assessed as ‘Medium’ (within 2 years) and sensitivity is therefore assessed as ‘Medium’.  This assessment is relatively precautionary and it should be noted the degree of resilience will be mediated by the character of the impact.  The recovery of small areas of surficial damage in thick reefs is likely to occur through tube repair and may be relatively rapid. 

Sabellaria spinulosa reef biotopes are directly exposed to physical damage that affects the surface layers (abrasion) and penetrates deeper beneath the surface of the reef.   No quantitative studies were found and although Vorberg (2000) is widely cited (see above in the abrasion section) the study used a light shrimp trawl on Sabellaria alveolata reefs and the relevance to Sabellaria spinulosa and the use of heavy fishing trawls is questionable.

Sabellaria spinulosa reefs in the Wadden Sea suffered great losses in the 1950s which are thought to be due to heavy anchor chains being trawled over grounds in association with shrimp fishing (Reise & Schubert, 1987; JNCC, 2013).  It is believed that local fishermen targeted areas of Sabellaria spinulosa reef due to their association with the brown shrimp Crangon crangon, and that deliberate attempts to remove the reefs were made so that fishing gear was not snagged and damaged  (Defra, 2004; JNCC, 2013).  Similar activity has been reported by fishermen at Ramsgate on Sabellaria spinulosa reefs in the Thames sea area but no direct evidence has been identified (Fariñas-Franco, 2014). 

Other studies have found significant evidence of trawl scars from unspecified fisheries through Sabellaria spinulosa reefs (Collins, 2003; Pearce et al., 2007) indicating that damage from fishing gear is a real possibility (Hendrick et al., 2011).  Obvious evidence of the destruction of Sabellaria spinulosa reef clumps by a beam trawler has been reported off the coast of Swanage, Dorset (Collins, 2003; cited from Benson et al., 2013).  The loss of reefs within a monitoring zone may have been due to bottom trawling based on the presence of trawl scars within the survey area, although the loss cannot be directly attributed to this activity based on the lack of direct observation (Pearce et al., 2011a).

Sabellaria spinulosa reefs remain extensive despite clear damage from bottom trawling at Hastings Shingle Bank (Cooper et al., 2007; Pearce et al., 2007) and at the Thanet offshore wind farm site (Pearce et al., in press).  However, in other areas such as the Wadden Sea (Riesen & Reise, 1982) and Morecambe Bay (see references in Holt et al., 1998), reefs which have been thought to have been trawled have disappeared and have not recovered.  It is acknowledged that the limited evidence available does not allow these losses to be directly attributed to fishing.

At the Hastings site, a newly developed reef (six months old) demonstrated the same multivariate community structure of fauna inhabiting a nearby reef that had been observed over the past 5 years (Pearce et al., 2007).  This suggests that the epifauna community associated with Sabellaria spinulosa reefs could also recover from fishing activity quickly, but it should be noted that the older reef had experienced on-going fishing activity and so the associated assemblage may be at a relatively early successional stage (Pearce et al., 2007). The quick recovery of the reef and associated biota was not seen in the Wadden Sea after shrimping activity in the 1950’s.  Instead, together with a loss of mussel beds and seagrass, community composition in the subtidal zone changed and a significant decline in sessile species was observed (Reise et al., 1989; Buhs & Reise, 1997; Reise & Buhs, 1999; Reise, 2005).

Sensitivity assessment. Structural damage to the seabed sub-surface is likely to damage and break-up tube aggregations leading to the loss of reef within the footprint of direct impact.  Sabellaria spinulosa is assessed as having a resistance of ‘None’ to this pressure (removal of >75% of the reef in the pressure footprint).  Based on evidence (Pearce et al., 2007; Pearce et al., 2011a) resilience was assessed as ‘Medium’, therefore, the sensitivity of Sabellaria spinulosa biotopes is considered to be ‘Medium’.  

Sabellaria spinulosa do not rely on light penetration for photosynthesis, it is also believed that visual perception is limited and that this species does not rely on sight to locate food or other resources.  In a recent review of the sensitivity of Sabellaria spinulosa reefs to anthropogenic disturbance,  Fariñas-Franco et al.  (2014) concluded that impacts on Sabellaria spinulosa due to a decrease in water clarity resulting from an increase in suspended solids (inorganic or organic) are unlikely although no thresholds regarding tolerance or intolerance were found.  Decreases in suspended particles that reduce the supply of food or tube-building materials may, however, negatively impact this species (Davies et al., 2009; Last et al., 2011).

Sabellaria spinulosa relies on a supply of suspended solids and organic matter in order to filter feed and build protective tubes and so they are often found in areas with high levels of turbidity.  Davies, et al. (2009) and Last et al. (2011) developed Vortex Resuspension Tanks (VoRT) which are able to test the effects of a change in the composition of suspended sediment on benthic species.  This laboratory experiment manipulated turbidity and current flow and demonstrated the susceptibility of Sabellaria spinulosa to a decrease in suspended particulate matter (SPM).  A clear erosion of tubes was observed in the absence of SPM and subsequent starvation of tube building materials.  At high and intermediate sediment regimes (high SPM ~71 mg / l) conditions were comparable to what might be expected within only a few hundred meters distance of a primary aggregate extraction site and Sabellaria spinulosa maintained a cumulative growth rate at these rates of SPM.  This supports the view that availability of suspended particles is necessary for Sabellaria spinulosa development and that tolerance of elevated levels is likely (Davies et al., 2009).

Indirect evidence for the tolerance of Sabellaria spinulosa for changes in turbidity is provided by the persistence of reefs on the outskirts of aggregate dredging areas (Pearce et al., 2007, 2011a) which appear unaffected by extraction which is likely to have led to sediment plumes.  Such plumes, however, are short-lived (Tillin et al., 2011) and, therefore, the long-term effect depends on the duration of dredging activities.

Tillin, (2010) used logistic regression to develop statistical models that indicate how the probability of occurrence of the congener Sabellaria alveolata changes over environmental gradients within the Severn Estuary.  The model predicted response surfaces were derived for each biotope for each of the selected habitat variables, using logistic regression.  From these response surfaces the optimum habitat range for each biotope could be defined based on the range of each environmental variable where the probability of occurrence, divided by the maximum probability of occurrence, is 0.75 or higher.  These results identify the range for each significant variable where the habitat is most likely to occur.  The modelled ranges should be interpreted with caution and apply to the Severn Estuary alone (which experiences large tidal ranges, high currents, and extremely high suspended sediment loads and is, therefore, distinct from many other estuarine systems).  However, these ranges do provide some useful information on environmental tolerances.  The models indicate that for subtidal Sabellaria alveolata the optimal mean neap sediment concentrations range from 515.7-906 mg/l and optimal mean spring sediment concentrations range from 855.3-1631 mg/l.  Although not directly applicable to S. spinulosa this data suggests that tube-building sabellariids are tolerant to very high levels of suspended sediment.  Fine sediments such as mud may clog the gills and feeding tentacles of some polychaetes and therefore, the potential impact will be mediated by the character of the pressure.

Sensitivity assessment. The benchmark for this pressure refers to a change in turbidity of one rank (see benchmark)  Sabellaria spinulosa do not photosynthesise and do not rely on sight to locate resources and, therefore, no effects are predicted for reef biotopes from an increase or decrease in clarity resulting from a change in one rank on the water framework directive scale.  Experiments (Davies et al., 2009) and predictive modelling (Tillin, 2010) indicate that tube building sabellariids can tolerate a broad range of suspended solids. Resistance to an increase or decrease at the pressure benchmark is therefore assessed as ‘High’ and resilience as ‘High’ (no impact to recover from). 

Sabellaria spinulosa are often found in areas of high water movement with some degree of sediment transport essential for tube-building and feeding.  Sabellaria spinulosa reefs adjacent to aggregate dredging areas appear unimpacted by dredging operations (Pearce et al., 2007; Pearce et al., 2011a).  Evidence suggests that given the dynamic sedimentary environments in which sabellariids live, their populations can certainly persevere in turbid conditions in spite of ‘typical’ natural levels of burial (Last et al., 2011) and that recovery from burial events is high.  

Direct evidence for the effects of siltation on Sabellaria spinulosa is limited to the experiments undertaken by Last et al., (2011).  Last et al. (2011) buried Sabellaria spinulosa worms (isolated into artificial tubes), under three different depths of sediment – shallow (2cm), medium (5cm) and deep (7cm).  The results indicate that Sabellaria spinulosa can survive short-term (32 days), periodic sand burial of up to 7 cm.  Last et al. (2011) suggested that the formation of ‘emergence tubes’ (newly created tubes extending to the surface) under sediment burial allowed Sabellaria spinulosa to tolerate gradual burial and that perhaps this mechanism allows for continued adult dispersal.  This mechanism occurred most rapidly throughout the 8-day burial at ~1 mm per day (Last et al., 2011) but even though tube-growth still seems possible under burial, it is likely that a dumping of fine and coarse material will block feeding apparatus and, therefore, worm development will be curtailed.

The evidence above suggests that Ross worm reefs are sensitive to damage from siltation events (Hendrick et al., 2011).  However, at the pressure benchmark, the depth of burial is likely to be similar or less than that experienced during natural storm events that move sediments and, in areas of high water movement, deposits of fine sediments are likely to be remobilised and moved.

Sensitivity assessment. In areas of high water flow dispersion of fine sediments may be rapid and this will mitigate the magnitude of this pressure by reducing the time exposed.  Based on the experiments by Last et al. (2011) which are considered relevant to the pressure benchmark, resistance and resilience are assessed as ‘High’ and this biotope is considered to be 'Not sensitive'.   

Sabellaria spinulosa reefs adjacent to aggregate dredging areas appear unimpacted by dredging operations (Pearce et al., 2007; Pearce et al., 2011a).  Evidence suggests that given the dynamic sedimentary environments in which sabellariids live, their populations can certainly persevere in turbid conditions in spite of ‘typical’ natural levels of burial (Last et al., 2011) and that recovery from burial events is high.  The congener S. alveolata was reported to survive short-term burial for days and even weeks in the south west as a result of storms that altered sand levels up to two meters, they were, however, killed by longer-term burial (Earll & Erwin 1983).

Direct evidence for the effects of siltation on Sabellaria spinulosa is limited to the experiments undertaken by Last et al. (2011).  The experimental conditions do not, however, relate to the pressure benchmark (30 cm of siltation in a single event).  Last et al., (2011) buried Sabellaria spinulosa worms (isolated into artificial tubes), under three different depths of sediment – shallow (2 cm), medium (5 cm) and deep (7 cm).  The results indicate that Sabellaria spinulosa can survive short-term (32 days), periodic sand burial of up to 7 cm.  Last et al. (2011) suggested that the formation of ‘emergence tubes’ (newly created tubes extending to the surface) under sediment burial allowed Sabellaria spinulosa to tolerate gradual burial and that perhaps this mechanism allows for continued adult dispersal.  This mechanism occurred most rapidly throughout the 8-day burial at ~1 mm per day (Last et al., 2011) but even though tube-growth still seems possible under burial, it is likely that a dumping of fine and coarse material will block feeding apparatus and therefore worm development will be curtailed.

A Sabellaria spinulosa reef off the coast of Dorset has shown periodic burial from large sand waves (Collins, 2003).  The displacement of some colonies that had established themselves on a gas pipeline 1 km off the coast of Aberdeen was also associated with burial (Mistakidis, 1956; cited by Holt et al., 1998).  Furthermore the loss of a 2 km² area of Ross worm reef in Jade Bay, North Sea was attributed to burial as a consequence of mud deposition, although fishing activity may have contributed to the decline (Dörjes, 1992, cited from Hendrick et al., 2011). 

The evidence above suggests that Sabellaria spinulosa reefs are sensitive to damage from siltation events (Hendrick et al., 2011).  However, recovery is likely to be rapid given that larval dispersal is not interrupted and new reefs are likely to be able to establish themselves over old buried ones as postulated by (Fariñas-Franco et al., 2014).

Sensitivity assessment. No direct evidence was found for the length of time that Sabellaria spinulosa can survive beneath 30 cm of sediment.  In areas of high water flow dispersion of fine sediments may be rapid and this will mitigate the magnitude of this pressure by reducing the time exposed.  However, this mitigating effect was not taken into account as it depends on site-specific conditions.  Resistance was assessed as ‘None’ due to the depth of overburden. Resilience was assessed as ‘Medium’ (2-10 years) and sensitivity was therefore categorised as ‘Medium’.  

Not assessed.

No evidence.

Not relevant.

Not relevant.

No direct evidence was found to assess this pressure. As the larvae of Sabellaria spinulosa are planktonic and are transported by water movements, barriers that reduce the degree of tidal excursion may alter the supply of Sabellaria spinulosa to suitable habitats from source populations. However, the presence of barriers may enhance local population supply by preventing the loss of larvae from enclosed habitats.  This species is therefore potentially sensitive to barriers that restrict water movements, whether this will lead to beneficial or negative effects will depend on whether enclosed populations are sources of larvae or are ‘sink’ populations that depend on outside supply of larvae to sustain the local population.

Sensitivity assessment. As this habitat is potentially sensitive to changes in tidal excursion and exchange, resistance is assessed as ‘Medium’ and resilience as ‘High’, sensitivity is, therefore ‘Low’. It should be noted that offshore circalittoral habitats are unlikely to be exposed to this pressure.

Not relevant’ to seabed habitats.  NB. Collision by grounding vessels is addressed under ‘surface abrasion’.

Not relevant.

Sabellaria spinulosa is not farmed or translocated and, therefore, this pressure is 'Not relevant' to this biotope.

No direct evidence relating to the impacts of the introduction of non-indigenous species on Sabellaria spinulosa reefs was found to support this assessment by Gibb et al. (2014). For many of the non-indigenous species that are found in UK seabed habitats, there are no records to suggest that their distribution overlaps with Sabellaria spinulosa reefs.

The oyster drill, Urosalpinx cinerea, is not known to predate on polychaetes (Brown & Richardson, 1988), therefore, their introduction is not considered a threat to Sabellaria spinulosa. There is, however, some overlap between the environmental niche of Sabellaria spinulosa and the oysters that Urosalpinx cinerea selectively feed on (Brown & Richardson, 1988).  Japanese wireweed Sargassum muticum and green sea fingers Codium fragile have the potential to compete for space where Sabellaria spinulosa reefs occur intertidally, however, intertidal biotopes are not included in this assessment and these species are unlikely to impact deeper subtidal reefs. No records of the carpet sea squirt Didemnum vexillum, the reef building serpulid Ficopomatus enigmaticus, the colonial sea squirt Perophora japonica, or Japanese kelp Undaria pinnatifida, suggest these species occur on or near Sabellaria spinulosa reefs. However, further spread may impact subtidal Sabellaria spinulosa reefs through smothering or competition, although this is entirely speculative.

Sabellaria spinulosa reefs support a variety of attached epifauna including species of bryozoans, hydroids and sponges. As Sabellaria spinulosa reefs are known to support encrusting organisms without apparent adverse effect, the stalked sea squirt Styela clava, as a solitary sea squirt is considered unlikely to have greater negative impacts than native species.

Two species that potentially pose a threat to Sabellaria spinulosa reefs are the Pacific oyster Magallana gigas and the slipper limpet Crepidula fornicata. Reefs of Sabellaria alveolata in the bay of Mont Saint Michel, France are becoming increasingly colonized by the Pacific oyster Magallana gigas (Dubois et al. 2006). Given the high filtration rates of Crassostrea gigas, it is believed that they can out-compete Sabellaria alveolata for feeding resources (Dubois et al. 2006). In the Wadden Sea, Magallana gigas has replaced blue mussels (Diederich, 2005; 2006) suggesting that Magallana gigas may impact filter feeding, reef forming organisms in general. The reasons underlying the species shift from Mytilus edulis to Magallana gigas have not been elucidated, however, and may be due to recent changes in climactic conditions (Thieltges, 2005) rather than competitive interactions. It should be noted that even though Magallana gigas is distributed throughout UK waters following an initial introduction in 1926 (Linke, 1951) there is currently no evidence, in the absence of any targeted studies, that this species is impacting native Sabellaria spinulosa or Sabellaria alveolata reefs (Crisp, 1964; Hendrick, et al. 2011).

When the slipper limpet Crepidula fornicata settles in an area it can increase the amount of pseudofaeces and subsequently the substratum may be altered from hard substratum to soft sediment which will reduce the substratum available for settlement by other species. This was observed when a 28-30% mortality of Mytilus edulis occurred after the introduction of Crepidula fornicata to mussel beds (Thieltges, 2005). Crepidula fornicata has been recorded in association with Sabellaria spinulosa reefs at Hastings Shingle Bank (up to 66 individuals per grab, Pearce, 2007) and in lower numbers in the East Coast REC area (maximum 4 per grab, Pearce et al., 2011a). The relationship between Crepidula fornicata and Sabellaria spinulosa has not been investigated. However, potential impacts on Sabellaria spinulosa reefs could occur through changes to substratum suitability or other interactions.

Sensitivity assessment. No evidence was found that non-indigenous species are currently significantly impacting Sabellaria spinulosa reef biotopes. Based on current evidence, resistance is therefore assessed as ‘High’ and resilience as ‘High’ (no impact to recover from), so that all the Sabellaria spinulosa reef biotopes are assessed as ‘Not Sensitive’. However, it should be noted that Crepidula fornicata and Magallana gigas may pose a potential threat in terms of competition for food and space and so this assessment may require updating in the future as the distributions and interactions between these species are better understood.

No evidence was found for adverse impacts of microbial pathogens on Sabellaria spinulosa.

Sabellaria spinulosa may be directly removed or damaged by static or mobile gears that are targeting other species. These direct, physical impacts are assessed through the abrasion and penetration of the seabed pressures. The sensitivity assessment for this pressure considers any biological effects resulting from the removal of target species on Sabellaria spinulosa biotopes.

Sabellaria spinulosa has no economic value and is not commercially harvested. Gibb et al. (2014) reviewed the evidence regarding potential biological effects of the removal of other target species on the Sabellaria spinulosa reef biotopes. Experimental laboratory work reported that scallop shells, especially Pecten maximus, induced Sabellaria spinulosa larvae to settle (Earll & Erwin 1983). However; the settlement-inducing property of Pecten maximus shells related mostly to the upper valve which was covered in sand grains (an existing requirement of larvae settlement) and given the diverse range of substrata that Sabellaria spinulosa have been reported in (see physical change pressure below) it is unlikely that the removal of scallops will have a significant negative impact on larval recruitment.

Gibb et al. (2014) suggest that the removal of target species that prey on Sabellaria spinulosa could potentially be beneficial to this species. Assessment of this indirect effect is limited by the lack of empirical evidence for predator-prey relationships. Sabellaria spinulosa reefs appear to be important nursery areas for commercially targeted flat fish including Dover sole (Bryony Pearce, pers comm). Stomach analysis of fish by Pearce (2011b) found that juvenile flatfish captured in reef areas including Dover sole, dab and plaice fed preferentially on Sabellaria spinulosa. Where these species are removed as target species (or as by-catch, given that commercial fisheries are unliklely to focus on juveniles), then predation rates on Sabellaria spinulosa could be reduced.  

Sensitivity assessment. Sabellaria spinulosa has no economic value and is not commercially harvested. The Sabellaria spinulosa biotopes are not, therefore directly impacted by this pressure and all biotopes within this group are considered, by default, to be ‘Not Sensitive’. The removal of target species that predate on Sabellaria spinulosa may have a potentially beneficial effect and the biotope is not considered to be sensitive to ecological effects resulting from their removal. 

Sabellaria spinulosa biotopes may be removed or damaged by static or mobile gears that are targeting other species. These direct, physical impacts are assessed through the abrasion and penetration of the seabed pressures. Evidence for ecological interactions between Sabellaria spinulosa and other species is limited. The removal of Sabellaria spinulosa predators as bycatch may be beneficial. Pearce et al. (2011b) found that as well as the commercially targeted species described for the removal of target species pressure, butterfish Pholis gunnelis and dragonet Callionymus lyra predated on Sabellaria spinulosa. Previous studies have also shown that Carcinus maenas feeds on Sabellaria spinulosa (Taylor 1962; Bamber & Irving, 1997). Other invertebrates such as Pandalus montagui and Asterias rubens found in association with Sabellaria spinulosa reefs may also be feeding on the worms or on species associated with the reefs rather than Sabellaria spinulosa. Due to the limited information available on predator-prey relationships, the impact of predator removal on Sabellaria spinulosa reef biotopes cannot be assessed.

Dense aggregations of the brittle star, Ophiothrix fragilis, have been suggested to compete with Sabellaria spinulosa for space and food and potentially to consume the gametes inhibiting recruitment (George & Warwick 1985). Removal of this species as by-catch could potentially be beneficial to the reef biotopes. A further potential interaction has been identified between Sabellaria spinulosa and the sand mason Lanice conchilega. It has been observed that sand stabilised by the sand mason Lanice conchilega is stable enough for colonization by Sabellaria alveolata (Larsonneur et al. 1994). It is believed that the same may also be possible for Sabellaria spinulosa, as Lanice conchilega and Sabellaria spinulosa are sometimes found together (Holt et al. 1997). However, a decline in Sabellaria spinulosa numbers was coincident with an increase in Lanice conchilega numbers (see Limpenny et al., 2010 and references therein). It is not clear from the available evidence therefore whether removal of Lanice conchilega would have any impact on Sabellaria spinulosa reefs (Foster-Smith 2001a).

Sensitivity assessment.The biogenic structure created by the Sabellaria spinulosa worms is the key characterizing feature of this biotope.  Removal of the worms and tubes as by-catch would remove the biotope and hence this group is considered to have a resistance of 'None' to this pressure and to have ‘Medium’ recovery. Sensitivity is, therefore ‘Medium’.