The Marine Life Information Network

Global warming (extreme)

Extreme emission scenario (by the end of this century 2081-2100) benchmark of:

• A 5°C rise in SST and NBT (coastal to the shelf seas),

• A 6°C rise in surface air temperature (in eulittoral and supralittoral habitats).

• A 1°C rise in Deep-sea habitats (>200 m) off the continental shelf, and

• A 5°C rise in surface air temperature in intertidal habitats exclusive to Scotland (Global warming pressure definitions).

Evidence

The honeycomb worm, Sabellaria alveolata, is a warm-temperate species and is distributed from Scotland to Morocco, (Gruet, 1986, Firth et al., 2015) including extensive reefs in the Mediterranean (La Porta & Nicoletti, 2009). This suggests that this species will be tolerant of increases in temperature.

This appears to be backed up by in situ observations of this species. Studies at Hinkley Point, Somerset, found that growth of the tubes in the winter was considerably greater in the cooling water outfall, where the water temperature was raised by around 8-10°C than at a control site, although the size of the individual worms themselves seemed to be unaffected (Bamber & Irving, 1997). Dubois et al. (2007) observed that in autumn where water temperatures are 8°C higher than in spring, a shorter period was required for larvae to metamorphose. The growth of Sabellaria alveolata is severely restricted below 5°C (Gruet, 1982, cited in Holt et al., 1998). Cunningham et al. (1984) reported increasing growth rates with temperatures up to 20°C. Long-term data for the coast of northern England and Wales shows that Sabellaria alveolata has colonized new locations, recolonizing areas from where it disappeared after the cold winter of 1962/63, and increased its abundance at many locations (Firth et al., 2015). This is possibly in response to recent warming (Firth et al., 2015).

Different spawning regimes, which may be due to different water temperatures, have been observed where conditions for a more northern population are less favourable and led to single annual spawning events of shorter duration (Culloty et al., 2010). Populations of Sabellaria alveolata in Morocco and the Mediterranean may be close to their upper thermal limit, as Muir et al., 2016 found that when individuals were exposed to 25°C, changes in lipid composition appeared to show a stress response.

Sensitivity assessment. Under the middle and high emission and extreme scenarios, seawater temperatures are expected to rise by 3-5°C to potential southern summer temperatures of 22-24°C and northern summer temperatures of 17-19°C. As this species is intertidal, it would also be affected by an increase in air temperature. Currently, summer temperatures can reach up to an average of 25°C, although the highest temperature recorded in 1961-2010 was 38.5°C (Perry & Golding, 2011). If air temperatures were to rise by 3, 4, or 6°C by the end of the century (middle, high and extreme emission scenarios, respectively), this could lead to temperatures reaching average summer high temperatures of between 28 - 32°C.

There is no experimental evidence of the impact of ocean warming on this species but biogeographic distribution is often a good predictor of temperature tolerance (Jeffree & Jeffree, 1994). Sabellaria alveolata is known to form extensive reefs in the Tyrrhenian Sea where summer water temperatures reach 27°C (www.seatemperature.org), although in the Mediterranean this species often occurs in the shallow subtidal (La Porta & Nicoletti, 2009).  This species also occurs in Morocco, where air temperatures can regularly reach 28°C in the summer months (www.weather-and-climate.com). In Scotland, this species is at the northern-most edge of its distribution and increases in seawater temperature are likely to be beneficial. This preference for warmer waters is backed up by observations of increased growth at of this species in the cooling waters of the power station at Hinkley Point in Somerset (Bamber & Irving, 1997). 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.

Global warming (high)

High emission scenario (by the end of this century 2081-2100) benchmark of:

• A 4°C rise in SST, NBT (coastal to the shelf seas) and surface air temperature (in eulittoral and supralittoral habitats).

• A 1°C rise in Deep-sea habitats (>200 m) off the continental shelf, and

• A 3°C rise in surface air temperature in intertidal habitats exclusive to Scotland. 

Evidence

The honeycomb worm, Sabellaria alveolata, is a warm-temperate species and is distributed from Scotland to Morocco, (Gruet, 1986, Firth et al., 2015) including extensive reefs in the Mediterranean (La Porta & Nicoletti, 2009). This suggests that this species will be tolerant of increases in temperature.

This appears to be backed up by in situ observations of this species. Studies at Hinkley Point, Somerset, found that growth of the tubes in the winter was considerably greater in the cooling water outfall, where the water temperature was raised by around 8-10°C than at a control site, although the size of the individual worms themselves seemed to be unaffected (Bamber & Irving, 1997). Dubois et al. (2007) observed that in autumn where water temperatures are 8°C higher than in spring, a shorter period was required for larvae to metamorphose. The growth of Sabellaria alveolata is severely restricted below 5°C (Gruet, 1982, cited in Holt et al., 1998). Cunningham et al. (1984) reported increasing growth rates with temperatures up to 20°C. Long-term data for the coast of northern England and Wales shows that Sabellaria alveolata has colonized new locations, recolonizing areas from where it disappeared after the cold winter of 1962/63, and increased its abundance at many locations (Firth et al., 2015). This is possibly in response to recent warming (Firth et al., 2015).

Different spawning regimes, which may be due to different water temperatures, have been observed where conditions for a more northern population are less favourable and led to single annual spawning events of shorter duration (Culloty et al., 2010). Populations of Sabellaria alveolata in Morocco and the Mediterranean may be close to their upper thermal limit, as Muir et al., 2016 found that when individuals were exposed to 25°C, changes in lipid composition appeared to show a stress response.

Sensitivity assessment. Under the middle and high emission and extreme scenarios, seawater temperatures are expected to rise by 3-5°C to potential southern summer temperatures of 22-24°C and northern summer temperatures of 17-19°C. As this species is intertidal, it would also be affected by an increase in air temperature. Currently, summer temperatures can reach up to an average of 25°C, although the highest temperature recorded in 1961-2010 was 38.5°C (Perry & Golding, 2011). If air temperatures were to rise by 3, 4, or 6°C by the end of the century (middle, high and extreme emission scenarios, respectively), this could lead to temperatures reaching average summer high temperatures of between 28 - 32°C.

There is no experimental evidence of the impact of ocean warming on this species but biogeographic distribution is often a good predictor of temperature tolerance (Jeffree & Jeffree, 1994). Sabellaria alveolata is known to form extensive reefs in the Tyrrhenian Sea where summer water temperatures reach 27°C (www.seatemperature.org), although in the Mediterranean this species often occurs in the shallow subtidal (La Porta & Nicoletti, 2009).  This species also occurs in Morocco, where air temperatures can regularly reach 28°C in the summer months (www.weather-and-climate.com). In Scotland, this species is at the northern-most edge of its distribution and increases in seawater temperature are likely to be beneficial. This preference for warmer waters is backed up by observations of increased growth at of this species in the cooling waters of the power station at Hinkley Point in Somerset (Bamber & Irving, 1997). 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.

Global warming (middle)

Middle emission scenario (by the end of this century 2081-2100) benchmark of:

• A 3°C rise in SST, NBT (coastal to the shelf seas) and surface air temperature (in eulittoral and supralittoral habitats).

• A 1°C rise in Deep-sea habitats (>200 m) off the continental shelf.

• A 2°C rise in surface air temperature in intertidal habitats exclusive to Scotland. 

Evidence

The honeycomb worm, Sabellaria alveolata, is a warm-temperate species and is distributed from Scotland to Morocco, (Gruet, 1986, Firth et al., 2015) including extensive reefs in the Mediterranean (La Porta & Nicoletti, 2009). This suggests that this species will be tolerant of increases in temperature.

This appears to be backed up by in situ observations of this species. Studies at Hinkley Point, Somerset, found that growth of the tubes in the winter was considerably greater in the cooling water outfall, where the water temperature was raised by around 8-10°C than at a control site, although the size of the individual worms themselves seemed to be unaffected (Bamber & Irving, 1997). Dubois et al. (2007) observed that in autumn where water temperatures are 8°C higher than in spring, a shorter period was required for larvae to metamorphose. The growth of Sabellaria alveolata is severely restricted below 5°C (Gruet, 1982, cited in Holt et al., 1998). Cunningham et al. (1984) reported increasing growth rates with temperatures up to 20°C. Long-term data for the coast of northern England and Wales shows that Sabellaria alveolata has colonized new locations, recolonizing areas from where it disappeared after the cold winter of 1962/63, and increased its abundance at many locations (Firth et al., 2015). This is possibly in response to recent warming (Firth et al., 2015).

Different spawning regimes, which may be due to different water temperatures, have been observed where conditions for a more northern population are less favourable and led to single annual spawning events of shorter duration (Culloty et al., 2010). Populations of Sabellaria alveolata in Morocco and the Mediterranean may be close to their upper thermal limit, as Muir et al., 2016 found that when individuals were exposed to 25°C, changes in lipid composition appeared to show a stress response.

Sensitivity assessment. Under the middle and high emission and extreme scenarios, seawater temperatures are expected to rise by 3-5°C to potential southern summer temperatures of 22-24°C and northern summer temperatures of 17-19°C. As this species is intertidal, it would also be affected by an increase in air temperature. Currently, summer temperatures can reach up to an average of 25°C, although the highest temperature recorded in 1961-2010 was 38.5°C (Perry & Golding, 2011). If air temperatures were to rise by 3, 4, or 6°C by the end of the century (middle, high and extreme emission scenarios, respectively), this could lead to temperatures reaching average summer high temperatures of between 28 - 32°C.

There is no experimental evidence of the impact of ocean warming on this species but biogeographic distribution is often a good predictor of temperature tolerance (Jeffree & Jeffree, 1994). Sabellaria alveolata is known to form extensive reefs in the Tyrrhenian Sea where summer water temperatures reach 27°C (www.seatemperature.org), although in the Mediterranean this species often occurs in the shallow subtidal (La Porta & Nicoletti, 2009).  This species also occurs in Morocco, where air temperatures can regularly reach 28°C in the summer months (www.weather-and-climate.com). In Scotland, this species is at the northern-most edge of its distribution and increases in seawater temperature are likely to be beneficial. This preference for warmer waters is backed up by observations of increased growth at of this species in the cooling waters of the power station at Hinkley Point in Somerset (Bamber & Irving, 1997). 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 (high)

High emission scenario benchmark: A marine heatwave occurring every two years, with a mean duration of 120 days, and a maximum intensity of 3.5°C (Marine heatwave pressure definitions).

Evidence

Marine heatwaves due to increased air-sea heat flux are predicted to occur more frequently and increase in duration and 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 alveolata, this species appears to 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 potentially suggested a stress response. Whilst high temperatures may cause stress, there is no evidence of mortality at seawater temperatures of 25°C. For example, when summer temperatures were increased from 18°C to 23°C for 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. A decrease in occupancy was only observed for the Bay of Biscay population, which the authors attributed to the poor state of the reef.

Curd et al. (2021) demonstrated that Sabellaria alveolata at sites within the UK (poleward) had smaller and more irregular shaped eggs in the winter, compared to worms closer to the equator. This variability was shown to be associated with several environmental drivers, including marine heatwaves, which exerted strong influence. Therefore, while long-term increases in SST may positivity impact the quality of gametes produced by this species in its northern range, extreme temperature increases exerted by marine heatwaves may decrease reproductive success.

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, and air temperatures exceeding 30°C across much of the UK.

Sabellaria alveolata is known to form extensive reefs in the Tyrrhenian Sea (La Porta & Nicoletti, 2009), where summer water temperatures reach 27°C (www.seatemperature.org) (see Global Warming). As this biotope occurs in the intertidal, this species will not only experience increased sea surface temperatures but will experience extreme air temperature increases also. In the Mediterranean, it can occur in the shallow subtidal (La Porta & Nicoletti, 2009) which may protect it from excessive air temperature increases, although this species also occurs in Morocco, where air temperatures can regularly reach 28°C in the summer months (www.weather-and-climate.com). As Sabellaria alveolata occurs on wave exposed coastlines on the mid and lower shore, wave splash may play an important role in preventing desiccation during emersion. 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, resistance has been assessed as ‘High’. As no recovery is likely necessary, resilience has been assessed as ‘High’, leading to an assessment of ‘Not sensitive’ for this biotope.

Marine heatwaves (middle)

Middle emission scenario benchmark:  A marine heatwave occurring every three years, with a mean duration of 80 days, with a maximum intensity of 2°C. 

Evidence

Marine heatwaves due to increased air-sea heat flux are predicted to occur more frequently and increase in duration and 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 alveolata, this species appears to 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 potentially suggested a stress response. Whilst high temperatures may cause stress, there is no evidence of mortality at seawater temperatures of 25°C. For example, when summer temperatures were increased from 18°C to 23°C for 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. A decrease in occupancy was only observed for the Bay of Biscay population, which the authors attributed to the poor state of the reef.

Curd et al. (2021) demonstrated that Sabellaria alveolata at sites within the UK (poleward) had smaller and more irregular shaped eggs in the winter, compared to worms closer to the equator. This variability was shown to be associated with several environmental drivers, including marine heatwaves, which exerted strong influence. Therefore, while long-term increases in SST may positivity impact the quality of gametes produced by this species in its northern range, extreme temperature increases exerted by marine heatwaves may decrease reproductive success.

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, and air temperatures exceeding 30°C across much of the UK.

Sabellaria alveolata is known to form extensive reefs in the Tyrrhenian Sea (La Porta & Nicoletti, 2009), where summer water temperatures reach 27°C (www.seatemperature.org) (see Global Warming). As this biotope occurs in the intertidal, this species will not only experience increased sea surface temperatures but will experience extreme air temperature increases also. In the Mediterranean, it can occur in the shallow subtidal (La Porta & Nicoletti, 2009) which may protect it from excessive air temperature increases, although this species also occurs in Morocco, where air temperatures can regularly reach 28°C in the summer months (www.weather-and-climate.com). As Sabellaria alveolata occurs on wave exposed coastlines on the mid and lower shore, wave splash may play an important role in preventing desiccation during emersion. 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, resistance has been assessed as ‘High’. As no recovery is likely necessary, resilience has been assessed as ‘High’, leading to an assessment of ‘Not sensitive’ for this biotope.

Ocean acidification (high)

High emission scenario benchmark: a further decrease in pH of 0.35 (annual mean) and corresponding 120% increase in H+ ions, seasonal aragonite saturation of 20% of UK coastal waters and North Sea bottom waters, and the aragonite saturation horizon in the NE Atlantic, off the continental shelf, occurring at a depth of 400 m by the end of this century 2081-2100 (Ocean acidification pressure definitions).

Evidence

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 alveolata or any species from the family Sabellariidae. Unlike the tube-dwelling, calcifying polychaetes from the family Serpulidae, Sabellaria alveolata does not form its tubes by calcification. Tubes are formed from coarse sand and shell grains cemented together with an adhesive protein cement secreted by the worm (Becker et al., 2012).

Non-calcifying polychaetes are 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 is variable among seasons and between females, but Sabellaria alveolata exhibits high fecundity, releasing an average of 100,000 eggs/individuals into the water column during a spawning cycle (Dubois, 2003).

Sensitivity Assessment. Direct evidence of the impact of ocean acidification on Sabellaria alveolata is lacking. However, non-calcifying polychaetes appear to be tolerant of changes in pH. 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.

Ocean acidification (middle)

Middle emission scenario benchmark: a further decrease in pH of 0.15 (annual mean) and a corresponding 35% increase in H+ ions with no coastal aragonite undersaturation and the aragonite saturation horizon in the NE Atlantic, off the continental shelf, at a depth of 800 m by the end of this century, 2081-2100. 

Evidence

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 alveolata or any species from the family Sabellariidae. Unlike the tube-dwelling, calcifying polychaetes from the family Serpulidae, Sabellaria alveolata does not form its tubes by calcification. Tubes are formed from coarse sand and shell grains cemented together with an adhesive protein cement secreted by the worm (Becker et al., 2012).

Non-calcifying polychaetes are 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 is variable among seasons and between females, but Sabellaria alveolata exhibits high fecundity, releasing an average of 100,000 eggs/individuals into the water column during a spawning cycle (Dubois, 2003).

Sensitivity Assessment. Direct evidence of the impact of ocean acidification on Sabellaria alveolata is lacking. However, non-calcifying polychaetes appear to be tolerant of changes in pH. 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 (extreme)

Extreme scenario benchmark: a 107 cm rise in average UK sea-level by the end of this century (2018-2100) (Sea-level rise pressure definitions).

Evidence

Sea-level rise is occurring through a combination of thermal expansion and ice melt. Sea levels have risen 1 to 3 mm/yr in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). Sea-level rise is expected to lead to substantial loss of intertidal habitats. Rocky shores backed by cliffs constitute about 80% of oceanic coastlines globally and in Britain, 42% of the coastline is hard rock, with many areas having cliffs behind the shore (Jackson & McIlvenny, 2011).

Jackson & McIlvenny (2011) predicted that under a 30 cm sea-level rise, between 10 and 27% of the extent of intertidal rocky shores in Scotland would be lost, whilst under a 190 cm of sea-level rise, between 26 - 50% would be lost. Using a modelling-based approach, Kaplanis et al. (2019) found that in San Diego County, loss of intertidal habitat would be most extreme within the first metre of sea-level rise, with 29.9% of intertidal rocky shore lost as a result of 20 cm sea-level rise, and 77.7% as a result of 100 cm of sea-level rise.

In the UK, intertidal Sabellaria alveolata beds occur on the mid and lower shore of exposed rocky coastlines. Wave action is an important driver of habitat quality for Sabellaria alveolata, supporting reef development by resuspending and transporting suitable sediment particles (Cunningham et al., 1984). High densities of Sabellaria alveolata are found on shores exposed to wave action (Anadόn, 1981; Dias & Paula, 2001), although reefs are generally absent from very exposed peninsulas such as the Lleyn, Pembrokeshire, and the extreme south-west of Cornwall, which probably relates to the effect of water movement on recruitment (Cunningham et al., 1984, cited from Holt et al., 1998). Habitat suitability modelling around Ireland determined that optimum tidal amplitude for Sabellaria alveolata is between 1.8 to 2.7 m (Firth et al., 2021). Understanding of 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 storms 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. It is difficult to assess the effect of sea-level rise scenarios on wave exposure or tidal energy as evidence predicts that any changes will be site-specific, therefore this aspect cannot be assessed. As this biotope occurs on the lower and mid-shore of the intertidal zone in the UK and is not in the subtidal, it is likely that an increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope. Beds may be able to expand their range and migrate upwards to compensate for sea-level rise, if not constrained by lack of suitable habitat (IPCC, 2019). If landward migration is not possible, it is expected that depth distribution of Sabellaria alveolata beds on littoral sediment will shrink in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery. In this assessment we have assessed on a worst-case-scenario basis, assuming that landward migration is not possible.

The mean tidal range in the UK varies from 127 cm in the Shetland Islands to 972 cm at Avonmouth, in the Bristol Channel (Woodworth et al., 1991). This large difference in tidal amplitudes suggests that this biotope will be more affected in some parts of the UK than others. In Scotland and Ireland, where mean tidal range is generally less than 3 m (Woodworth et al., 1991), more than half of this biotope may be lost under the extreme scenario, whereas in the Bristol Channel, where mean tidal range exceeds 9 m (Woodworth et al., 1991), only a small portion of this biotope may be lost. Under the medium emission scenario, resistance has been assessed as ‘Medium’, as it is likely that less than 25% of this biotope will be lost. Resilience has been assessed as ‘Very low’, due to the long-term nature of sea-level rise. Therefore, sensitivity is assessed as ‘Medium’.  Under the high emission and extreme scenarios, resistance has been assessed as ‘Low’, as more than 25% of this biotope could be lost. Resilience has been assessed as ‘Very low’, due to the long-term nature of sea-level rise. Therefore, sensitivity is assessed as ‘High’. 

Sea level rise (high)

High emission scenario benchmark: a 70 cm rise in average UK sea-level by the end of this century (2018-2100). 

Evidence

Sea-level rise is occurring through a combination of thermal expansion and ice melt. Sea levels have risen 1 to 3 mm/yr in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). Sea-level rise is expected to lead to substantial loss of intertidal habitats. Rocky shores backed by cliffs constitute about 80% of oceanic coastlines globally and in Britain, 42% of the coastline is hard rock, with many areas having cliffs behind the shore (Jackson & McIlvenny, 2011).

Jackson & McIlvenny (2011) predicted that under a 30 cm sea-level rise, between 10 and 27% of the extent of intertidal rocky shores in Scotland would be lost, whilst under a 190 cm of sea-level rise, between 26 - 50% would be lost. Using a modelling-based approach, Kaplanis et al. (2019) found that in San Diego County, loss of intertidal habitat would be most extreme within the first metre of sea-level rise, with 29.9% of intertidal rocky shore lost as a result of 20 cm sea-level rise, and 77.7% as a result of 100 cm of sea-level rise.

In the UK, intertidal Sabellaria alveolata beds occur on the mid and lower shore of exposed rocky coastlines. Wave action is an important driver of habitat quality for Sabellaria alveolata, supporting reef development by resuspending and transporting suitable sediment particles (Cunningham et al., 1984). High densities of Sabellaria alveolata are found on shores exposed to wave action (Anadόn, 1981; Dias & Paula, 2001), although reefs are generally absent from very exposed peninsulas such as the Lleyn, Pembrokeshire, and the extreme south-west of Cornwall, which probably relates to the effect of water movement on recruitment (Cunningham et al., 1984, cited from Holt et al., 1998). Habitat suitability modelling around Ireland determined that optimum tidal amplitude for Sabellaria alveolata is between 1.8 to 2.7 m (Firth et al., 2021). Understanding of 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 storms 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. It is difficult to assess the effect of sea-level rise scenarios on wave exposure or tidal energy as evidence predicts that any changes will be site-specific, therefore this aspect cannot be assessed. As this biotope occurs on the lower and mid-shore of the intertidal zone in the UK and is not in the subtidal, it is likely that an increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope. Beds may be able to expand their range and migrate upwards to compensate for sea-level rise, if not constrained by lack of suitable habitat (IPCC, 2019). If landward migration is not possible, it is expected that depth distribution of Sabellaria alveolata beds on littoral sediment will shrink in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery. In this assessment we have assessed on a worst-case-scenario basis, assuming that landward migration is not possible.

The mean tidal range in the UK varies from 127 cm in the Shetland Islands to 972 cm at Avonmouth, in the Bristol Channel (Woodworth et al., 1991). This large difference in tidal amplitudes suggests that this biotope will be more affected in some parts of the UK than others. In Scotland and Ireland, where mean tidal range is generally less than 3 m (Woodworth et al., 1991), more than half of this biotope may be lost under the extreme scenario, whereas in the Bristol Channel, where mean tidal range exceeds 9 m (Woodworth et al., 1991), only a small portion of this biotope may be lost. Under the medium emission scenario, resistance has been assessed as ‘Medium’, as it is likely that less than 25% of this biotope will be lost. Resilience has been assessed as ‘Very low’, due to the long-term nature of sea-level rise. Therefore, sensitivity is assessed as ‘Medium’.  Under the high emission and extreme scenarios, resistance has been assessed as ‘Low’, as more than 25% of this biotope could be lost. Resilience has been assessed as ‘Very low’, due to the long-term nature of sea-level rise. Therefore, sensitivity is assessed as ‘High’. 

Sea level rise (middle)

Middle emission scenario benchmark: a 50 cm rise in average UK sea-level by the end of this century (2081-2100).

Evidence

Sea-level rise is occurring through a combination of thermal expansion and ice melt. Sea levels have risen 1 to 3 mm/yr in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). Sea-level rise is expected to lead to substantial loss of intertidal habitats. Rocky shores backed by cliffs constitute about 80% of oceanic coastlines globally and in Britain, 42% of the coastline is hard rock, with many areas having cliffs behind the shore (Jackson & McIlvenny, 2011).

Jackson & McIlvenny (2011) predicted that under a 30 cm sea-level rise, between 10 and 27% of the extent of intertidal rocky shores in Scotland would be lost, whilst under a 190 cm of sea-level rise, between 26 - 50% would be lost. Using a modelling-based approach, Kaplanis et al. (2019) found that in San Diego County, loss of intertidal habitat would be most extreme within the first metre of sea-level rise, with 29.9% of intertidal rocky shore lost as a result of 20 cm sea-level rise, and 77.7% as a result of 100 cm of sea-level rise.

In the UK, intertidal Sabellaria alveolata beds occur on the mid and lower shore of exposed rocky coastlines. Wave action is an important driver of habitat quality for Sabellaria alveolata, supporting reef development by resuspending and transporting suitable sediment particles (Cunningham et al., 1984). High densities of Sabellaria alveolata are found on shores exposed to wave action (Anadόn, 1981; Dias & Paula, 2001), although reefs are generally absent from very exposed peninsulas such as the Lleyn, Pembrokeshire, and the extreme south-west of Cornwall, which probably relates to the effect of water movement on recruitment (Cunningham et al., 1984, cited from Holt et al., 1998). Habitat suitability modelling around Ireland determined that optimum tidal amplitude for Sabellaria alveolata is between 1.8 to 2.7 m (Firth et al., 2021). Understanding of 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 storms 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. It is difficult to assess the effect of sea-level rise scenarios on wave exposure or tidal energy as evidence predicts that any changes will be site-specific, therefore this aspect cannot be assessed. As this biotope occurs on the lower and mid-shore of the intertidal zone in the UK and is not in the subtidal, it is likely that an increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope. Beds may be able to expand their range and migrate upwards to compensate for sea-level rise, if not constrained by lack of suitable habitat (IPCC, 2019). If landward migration is not possible, it is expected that depth distribution of Sabellaria alveolata beds on littoral sediment will shrink in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery. In this assessment we have assessed on a worst-case-scenario basis, assuming that landward migration is not possible.

The mean tidal range in the UK varies from 127 cm in the Shetland Islands to 972 cm at Avonmouth, in the Bristol Channel (Woodworth et al., 1991). This large difference in tidal amplitudes suggests that this biotope will be more affected in some parts of the UK than others. In Scotland and Ireland, where mean tidal range is generally less than 3 m (Woodworth et al., 1991), more than half of this biotope may be lost under the extreme scenario, whereas in the Bristol Channel, where mean tidal range exceeds 9 m (Woodworth et al., 1991), only a small portion of this biotope may be lost. Under the medium emission scenario, resistance has been assessed as ‘Medium’, as it is likely that less than 25% of this biotope will be lost. Resilience has been assessed as ‘Very low’, due to the long-term nature of sea-level rise. Therefore, sensitivity is assessed as ‘Medium’.  Under the high emission and extreme scenarios, resistance has been assessed as ‘Low’, as more than 25% of this biotope could be lost. Resilience has been assessed as ‘Very low’, due to the long-term nature of sea-level rise. Therefore, sensitivity is assessed as ‘High’. 

Temperature increase (local)

Benchmark. A 5°C increase in temperature for one month, or 2°C for one year (Temperature change pressure definition).

Evidence

Sabellaria alveolata is a southern species reaching their northern limit in Britain and Ireland, whose global distribution extends south to Morocco (Gruet, 1982; Firth et al., 2015; Curd et al., 2023), with reports of occurrences as far south as the Bay of Bengal (Pradhan et al., 2025). Studies at Hinkley Point, Somerset, found that growth of the tubes in the winter was considerably greater in the cooling water outfall, where the water temperature was raised by around 8 to 10°C than at a control site, although the size of the individual worms themselves seemed to be unaffected (Bamber & Irving, 1997).

Curd et al. (2021) demonstrated the warm-adapted nature of this species by showing that individuals closer to the equator were less physiologically stressed and produced fewer irregularly shaped eggs in the winter compared to those towards the poles. Therefore, increases in SST may facilitate the physiological functioning and reproductive success of Sabellaria alveolata in the more northern parts of their range. Dubois et al. (2007) observed that in autumn where water temperatures are 8°C higher than in spring, a shorter period was required for larvae to metamorphose. Spawning regimes may differ due to different water temperatures. Where conditions for a more northern population are less favourable, they lead to single annual spawning events of shorter duration (Culloty et al., 2010).

Intertidal populations of Sabellaria alveolata are susceptible to low temperatures in winter (Crisp, 1964; Firth et al., 2015, 2021a). In north Wales, increases in Sabellaria alveolata abundance and colonization of previously uninhabited sites were reported during a period of warming from the mid-1980’s to the 2000s (Firth et al., 2015).

Modelling of habitat suitability has identified increases in SST as the second most influential variable in determining suitable habitat for Sabellaria alveolata (after wave action; Curd et al., 2023). Domy et al. (2023) modelled habitat suitability for Sabellaria alveolata around the UK given predicted SST increases of up to 3.2ºC. The results suggested that availability of highly suitable habitat could increase from 5.8% to 44% of the UK coastline. Their model suggested that most range expansion was expected to occur along the south and west coasts, potentially allowing for more connectivity between reefs. In addition, more suitable habitat was expected further north, particularly on the east coast of Scotland. However, specific niche requirements meant not all these available habitats would be reached, especially in the north, where hydrodynamic regimes such as enhanced currents, turbulent eddies and separated flows occur in the Hebrides (Domy et al., 2023). Range expansion may have been prevented in Ireland, where six regional sub-populations of Sabellaria alveolata have been identified. Three of these sub-populations align with persistent tidal fronts (Firth et al., 2021a). The boundary edges aligned with these fronts have been stable for the last 62 years, despite increases in SST that would predict range extension. Therefore, while the distribution of Sabellaria alveolata may otherwise increase with increases in SST, their expansion may be restricted by hydrographic regimes preventing colonization of more suitable habitat.

Sensitivity assessment. Based on distribution and temperature enhancement of duration and frequency of spawning, metamorphosis, growth rates and physiological functioning, Sabellaria alveolata is considered to be ‘Not sensitive’ to an increase in temperature at the pressure benchmark (therefore, resistance and resilience are both considered to be 'High').

Temperature decrease (local)

Benchmark. A 5°C decrease in temperature for one month, or 2°C for one year (Temperature change pressure definition).

Evidence

Sabellaria alveolata are a southern species reaching their northern limit in Britain and Ireland. Studies at Hinkley Point, Somerset, found that growth of the tubes in the winter was considerably greater in the cooling water outfall, where the water temperature was raised by around 8-10°C than at a control site, although the size of the individual worms themselves seemed to be unaffected (Bamber & Irving, 1997). 

Curd et al. (2021) concluded that this species was warm-adapted because individuals were more physiologically stressed at sites around the UK (poleward) compared to those towards the equator. Elevated levels of biomolecules indicative of thermal and oxidative stress, as well as those observed in energetically costly processes such as immune functioning were recorded for individuals at polewards sites. Whereas those in equatorward sites were least physiologically stressed. They also showed that worms in poleward sites had smaller and more irregular shaped eggs in the winter, compared to worms closer to the equator. This variability in reproductive traits was shown to be associated with several environmental drivers including extreme temperature variation in both air and seawater (i.e., cold spells and heatwaves) which exerted strong influence. Therefore, decreases in SST may further impede the physiological functioning and reproductive success of Sabellaria alveolata in the more northern parts of their range. Dubois et al. (2007) observed that in autumn where water temperatures are 8°C higher than in spring, a shorter period was required for larvae to metamorphose. Differences between spawning regimes which may be due to different water temperatures have been observed, where conditions for a more northern population are less favourable and lead to single annual spawning events of shorter duration (Culloty et al., 2010).

Intertidal populations of Sabellaria alveolata are susceptible to low temperatures in winter (Crisp, 1964) and modelling of habitat suitability determined that Sabellaria alveolata will likely disappear if minimum SST falls below 6°C (Firth et al., 2021a). Abundance of Sabellaria alveolata was shown to vary relative to SST. Along the coast of north Wales, Firth et al. (2015) demonstrated that an extreme cold spell in the winter of 1962/63 (mean winter SST of 5.3°C) caused declines in Sabellaria alveolata across north Wales and the Wirral, with complete mortality at two sites: Hilbre and Heysham. At Hilbre, recolonization and reef recovery were observed 40 years later. At Heysham, which recorded the coldest temperature during this winter (1.8°C), recovery took 20 years longer. After the same cold winter, Crisp (1964) reported immediate declines in Sabellaria alveolata abundance of up to 40% in Ireland and up to 95% at sites in Wales. Recovery of the site in Wales occurred by April 1963 from tube-building of surviving worms (Crisp, 1964). In addition, Firth et al. (2021a) identified four locations around the coast of Ireland where Sabellaria alveolata was present in the 1950’s, underwent localized extinctions in the 2000s, then recolonized and was either in recovery or fully recovered in the 2010’s. This temporal variability in Sabellaria abundance was also likely linked to the 1962/63 extreme cold winter (Firth et al., 2021a). In shorter timescales, other reefs around north Wales underwent localized extinctions of Sabellaria alveolata following a less severe cold spell in the winter of 2009/2010. At certain sites, recolonization of the site occurred within one year, however, at other sites, this species remained absent after three years (Firth et al., 2015).

Sensitivity assessment. The effects of acute decreases in temperature at the benchmark will depend on the seasonality of occurrence. Decreases in winter are likely to stress populations more than decreases in summer (although there may be effects on larval supply). At the centre of their UK range, adult Sabellaria alveolata are considered to have 'High' resistance to a chronic change at the pressure benchmark in summer. In extreme winters, resistance can be ‘Low’ or ‘None’, especially within the northern extreme of their range. Some previously abundant reefs fully recovered from extinction within one year. However, in other cases where larval supply and recruitment was limited or prevented by local hydrodynamics or temperature regimes (Firth et al., 2015, 2021a), recovery took several decades. Therefore, taken as a worst-case scenario, resistance is assessed as ‘None’ and resilience is ‘Very Low’. Therefore, sensitivity is assessed as ‘High’.

Salinity increase (local)

Benchmark. An increase in one MNCR salinity category above the usual range of the biotope or habitat (Salinity regime change pressure definition).

Evidence

Models of habitat suitability identified salinity as the third most influential variable determining the distribution of Sabellaria alveolata (Curd et al., 2023), supporting the idea that this biotope is restricted to areas of full salinity, defined as 30 to 35 ppt (Connor et al., 2004). The pressure benchmark of an increase in salinity is, therefore, 'Not relevant' to this biotope. However, it should be noted that reefs could be sensitive to hypersaline conditions above this benchmark. 

No empirical evidence was found to assess the impact of increases in salinity on adult, reef-forming populations. Quintino et al. (2008) examined through laboratory experiments the sub-lethal endpoints of brine exposure on Sabellaria alveolata larvae. Natural seawater where salinities had been increased using commercial salts used to prepare artificial seawater were used as the control. At a salinity of 36 (natural seawater artificially concentrated), 20% of Sabellaria alveolata developed abnormally. At a salinity of 40, this increased to about 70% of the larvae developed abnormally, clearly indicating the effect of increasing salinity on larvae. Although not directly relevant to the pressure benchmark, the experiments do suggest that increasing salinity would lead to lethal effects on larvae. It is not clear how these supply effects would ramify at the population level. Recruitment success varies between years (see resilience information) and a shortfall in one year may be compensated in another year when salinity returns to normal, providing the source population is unaffected.

Sensitivity assessment. The evidence above suggests that hypersaline conditions could temporarily impact larval recruitment but there is no evidence of the effects of hypersaline conditions on Sabellaria alveolata reefs or populations was found. Therefore, there is insufficient evidence to make an assessment.

Salinity decrease (local)

Benchmark. A decrease in one MNCR salinity category above the usual range of the biotope or habitat (Salinity regime change pressure definition detail).

Evidence

Models of habitat suitability have identified salinity as the third most influential variable determining the distribution of Sabellaria alveolata, having been recorded in salinities between 31.5 and 36.7 psu (Curd et al., 2023), supporting the idea that this biotope is restricted to areas of full salinity, defined as 30-35 ppt (Connor et al., 2004).

It is likely that Sabellaria alveolata can tolerate small declines in salinity as it occurs intertidally where freshwater inputs may lower salinity, either on a semi-permanent basis where rivers discharge into estuaries and bays, or where rainfall and land run-off cause an acute lowering of salinity. In the Bay of Mont-Saint-Michel, for example, where large reefs are found, salinities are lower (at <34.8) than in the open sea (Dubois et al., 2007). Although this biotope is reported to only occur in areas of full salinity (30-35 ppt), sublittoral reefs of Sabellaria alveolata are recorded in the Severn Estuary in areas of variable salinity (Connor et al., 2004). Lancaster (1993, cited from Holt et al., 1998) also found extensive, healthy hummocks of Sabellaria at Drigg, Cumbria, where there is a large freshwater input from the Drigg BNFL plant.

Sensitivity assessment. The evidence to assess this pressure is limited. Based on distribution with only occasional records within estuaries, this biotope is considered likely to be sensitive at the lower limits of the pressure benchmark (a change to reduced salinity; 18-30 ppt). Resistance is therefore assessed as ‘Low', as a reduction in salinity at the pressure benchmark is considered to result in the loss of most of the reef. Recovery from significant impacts can be much longer. Some abundant reefs can become extinct from a pressure and fully recover within one year, however, in other cases where larval supply and recruitment is limited or prevented by local hydrodynamics or temperature regimes (Firth et al., 2015, 2021a), recovery can take several decades. Therefore, where resistance is assessed as ‘Low’, especially within the northern extreme of their range, in the worst-case scenario resilience can be ‘Very Low’. Sensitivity is, therefore, ‘High’. The observed distribution of this biotope may be based on other factors than salinity, such as availability of suitable sediments, and confidence in this assessment is low. 

Water flow (tidal current) changes (local)

Benchmark. A change in peak mean spring bed flow velocity of between 0.1 m/s and 0.2 m/s for more than one year (Water flow pressure definition). 

Evidence

Holt et al. (1998) suggested that for Sabellaria alveolata reefs, the importance of currents vs waves in terms of sediment re-suspension and transport for tube-building varies regionally. In many British localities such as the south-west of England, much of Wales and the Cumbrian coast, waves seem more important, but in other areas such as parts of the Severn Estuary, tidal suspension is probably the key factor. Water flow in some areas will be a key driver of habitat suitability for Sabellaria alveolata due to the requirement for suspended sand for tube building and the supply of organic particles for food. Tests on the mechanical strength and properties of Sabellaria alveolata tubes were performed by Le Cam et al. (2011). These found that the biomineralized cement that the worms produce to bind sand grains into tubes confer wave resistance. Although thresholds of resistance are not known, the visco-elastic behaviour of the cement enables tubes to dissipate the mechanical energy of breaking waves and presumably also confers resistance to increased water flow rates (Le Cam et al. 2011).

Tillin (2010) used logistic regression to develop statistical models that indicate how the probability of occurrence of 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. 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 the results should be interpreted with caution, the modelled habitat suitability for Sabellaria alveolata indicated that the range of water flow tolerances is relatively broad. Furthermore, Firth et al. (2021a) modelled habitat suitability of Sabellaria alveolata around Ireland and determined that optimum tidal amplitude for this species fell between 1.8 and 2.7 m.

In general, sediment re-suspension and transport models indicate that sands are suspended by currents around 0.20-0.25 m/s and will stay in suspension until flow drops below 0.15-0.18 m/s (Wright et al., 2001). Sabellaria alveolata may be relatively insensitive to changes above these flow rates (although the upper tolerance limit is not clear). In sheltered habitats where the water flow rates are approaching the lower limits of water flow tolerance, a further reduction at the pressure benchmark may have negative impacts. Desroy et al. (2011) suggested that modifications to hydrodynamics (where current speed decreased downstream of new mussel farming infrastructure installations facing the reef) indirectly impacted sedimentary patterns and led to increased silt deposition resulting in the deterioration of Sabellaria alveolata reefs in the Bay of Mont-Saint-Michel, France.

Changes in water flow potentially have implications for larval transport and recruitment. Sabellaria alveolata is generally absent from very exposed peninsulas such as the Lleyn, Pembrokeshire, and the extreme south-west of Cornwall, which probably relates to the effect of water movement on recruitment (Cunningham et al., 1984, cited from Holt et al. 1998). However, behavioural responses by larvae to different flow rates may result in some control over movement. Dubois et al. (2007) observed the vertical migration of Sabellaria alveolata larvae during the tidal cycle, where larvae migrate upwards in the water column to faster near-surface currents and migrate down the water column on the ebb flow to where currents are weaker. This migration enhances landward transport of larvae to more suitable habitats and prevents seaward loss. Furthermore, larval transport may be hindered by local hydrodynamic regimes. In Ireland, six regional sub-populations of Sabellaria alveolata have been identified. three of which align with persistent tidal fronts (Firth et al., 2021a). The boundary edges aligned with these fronts have been stable for the last 62 years, despite increases in SST that would predict range extension. It is likely that these tidal fronts prevented the transport of larvae to suitable areas beyond.

Sensitivity assessment. 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 impacts, although Tillin (2010) modelled optimal flow speeds of 0.5 to 1.22 m/s. 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 optimal 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 (0.01 to 0.02 m/s), is not likely to result in mortality. Resistance is therefore assessed as ‘High’ and resilience as ‘High’ (no impact to recover from). This biotope is therefore considered to be ‘Not sensitive’.

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. (Emergence regime change pressure definition).

Evidence

A reduction in the amount of time spent under water could cause a proportion of exposed individual Sabellaria alveolata to die, as the worms can only feed when submerged. Sabellaria alveolata reefs also occur subtidally, a decrease in emergence time may have no physiological effect but may lead to increased predation on the reef. Furthermore, if exposed for more time, reefs may experience a higher trampling intensity from beach users. Cunningham et al. (1984) demonstrated that the reef recovered within 23 days from the effects of trampling (i.e. treading, walking or stamping on the reef structures) repairing minor damage to the worm tube porches. However, severe damage, estimated by kicking and jumping on the reef structure, resulted in large cracks between the tubes, and removal of sections (ca 15x15x10 cm) of the structure (Cunningham et al., 1984). 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. At another site, a smaller section (10x10x10 cm) was lost but after 23 days the space was already smaller due to rapid growth. In contrast, Plicanti et al. (2016) showed that even low intensity disturbance (being walked over once) caused significant damage to the reef by reducing the amount of intact bioconstructions. They demonstrated that two months after the disturbance, despite some increases, percentage cover of intact tubes had not recovered to control levels and remained significantly reduced at sites exposed to medium and high intensity trampling.

Modelling habitat suitability identified tidal amplitude as one of the most influential variables in determining suitable habitat for Sabellaria alveolata (Firth et al., 2021a). Around Ireland, the optimum tidal range for this species is between 1.8 to 2.7 m (Firth et al., 2021a). This study also noted that the occurrence of Sabellaria alveolata is favoured where the tidal amplitude is ‘moderately high’ between 2 and 2.5 m.

Sensitivity assessment. This biotope is recorded from the lower to middle shore. An increase in emergence is likely to reduce its upper limit and decrease the lower limit, and increase its exposure to trampling. Sabellaria alveolata has a preference for a moderately high tidal range (at least in Ireland, Firth et al., 2021a) so a decrease in emergence due to coastal constructions may be detrimental. Therefore, the biotope is probably sensitive to changes in emergence. Hence, resistance is assessed as ‘‘Low’ and recovery as ‘Medium’ (following habitat recovery). Sensitivity is, therefore assessed as ‘Medium’, but with 'Low' confidence

Wave exposure changes (local)

Benchmark. A change in near shore significant wave height of >3% but <5% for more than one year (Wave action pressure definition). 

Evidence

Models of habitat suitability have identified fetch and wave height as the some of the most influential variables in determining suitable habitat for Sabellaria alveolata (Firth et al., 2021a, Curd et al., 2023). Optimum wave exposure for these reefs was identified as moderate, such that wave height falls between 1.3 and 1.8 m (Firth et al., 2021a). Their models predicted that Sabellaria alveolata was unlikely to occur in areas where wave height exceeded 1.8 m (Firth et al., 2021a). Curd et al. (2023) suggested that there will be a loss in suitable habitat at the southerly range of this species, which will not be mitigated by a northern expansion, based on forecasted localised and regional climatologically induced changes to wave exposure, coupled with other environmental drivers. Wave action is an important driver of habitat quality for Sabellaria alveolata, supporting reef development by resuspending and transporting suitable sediment particles (Cunningham et al., 1984). High densities of Sabellaria alveolata are found on shores exposed to wave action (Anadόn, 1981; Dias & Paula, 2001), although reefs are generally absent from very exposed peninsulas such as the Lleyn, Pembrokeshire and the extreme south west of Cornwall, which probably relates to the effect of water movement on recruitment (Cunningham et al., 1984, cited from Holt et al., 1998).

Tests on the mechanical strength and properties of Sabellaria alveolata tubes were performed by Le Cam et al. (2011). These found that the biomineralized cement the worms produce to bind sand grains into tubes confer wave resistance. Although thresholds of resistance are not known, the visco-elastic behaviour of the cement enables tubes to dissipate the mechanical energy of breaking waves (Le Cam et al., 2011). A recent study in Italy investigated the effects of increased mechanical disturbance on the resistance of Sabellaria alveolata reef patches. Storari et al. (2024) simulated increased wave action by dropping a plastic bag filled with 10 L of water on the same reef patch three times over 15 months. The size of the reef patches was not reduced despite the extra mechanical disturbance. Rather, six months after the last disturbance, reef patches were on average 11% larger than the unmanipulated patches.

Wave action has also been linked to gamete quality in Sabellaria alveolata. Curd et al. (2021) showed that worms in poleward sites tended to be more physiologically stressed and had smaller and more irregular shaped eggs in the winter compared to worms closer to the equator. This variability in reproductive traits was shown to be associated with several environmental variables, including wave exposure, which is more variable at higher latitudes.

Sensitivity assessment. This biotope is recorded from moderately wave exposed and wave exposed habitats. A 3 to 5% change in significant wave height is small compared to the wave exposure it normally experiences. Firth et al. (2021a) reported that Sabellaria alveolata prefers a wave exposure between 1.3 and 1.8 m, but would be removed or excluded at > 1.8 m. However, the Sabellaria reefs are unlikely to be affected by a change at the benchmark level. Hence, resistance and resilience are assessed as ‘High’ and sensitivity as ‘Not sensitive’.

Transition elements & organo-metal contamination

Benchmark. Exposure of marine species or habitat to one or more relevant Transitional metal or organometal (e.g. TBT) contaminants via uncontrolled releases or incidental spills (Transitional metals and organometals pressure definition). 

Evidence

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

Hydrocarbon & PAH contamination

Benchmark. Exposure of marine species or habitat to one or more relevant hydrocarbon or polyaromatic hydrocarbon (PAH) contaminants via uncontrolled releases or incidental spills (Hydrocarbon & PAH pressure definition).

Evidence

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

Synthetic compound contamination

Benchmark. Exposure of marine species or habitat to one or more synthetic compound contaminants via uncontrolled releases or incidental spills (Synthetic compound contamination pressure definition).

Evidence

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

Radionuclide contamination

Benchmark. An increase in 10µGy/h above background levels (Radionuclides contamination pressure definition).

Evidence

Mauchline et al. (1964) examined concentration of radioactive isotopes by organisms on Windscale beach. Sabellaria alveolata built reefs with the smaller particles on the beach which adsorb the greatest amount of radioactivity per weight (due to surface-area effects). Thus Sabellaria reefs could concentrate radioactivity. However the study by Mauchline et al. (1964) did not look for or identify any potential negative effects on the worms such as changes in reproductive success or mortality rates.

Sensitivity assessment. No evidence.

Introduction of other substances

Benchmark. Exposure of marine species or habitat to one or more relevant "other" substances (solid, liquid or gas) contaminants via uncontrolled releases or incidental spills (Introduction of other substances pressure definition). 

Evidence

This pressure is Not assessed.

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) (deoxygenation pressure definition).

Evidence

No direct evidence was found to assess this pressure. As Sabellaria alveolata are primarily intertidal, respiration could occur during periods of emmersion so that this species is not exposed permanently to hypoxia/anoxia. This feature also occurs in relatively exposed areas on coarse substrates where water mixing is considered sufficient to prevent deoxygenation.

Sensitivity assessment. Based on habitat parameters mitigating this pressure, resistance is assessed as ‘Medium’ and recovery as ‘High’. Sensitivity is therefore assessed as ‘Low’.

Nutrient enrichment

Benchmark. Increased levels of the elements nitrogen, phosphorus, silicon, and iron in the marine environment compared to background concentrations (Nutrient enrichment pressure definition).

Evidence

Bertocci et al. (2017) assessed the effects of nutrient enrichment of Sabellaria alveolata in northern Portugal. Enrichment was performed by deploying 200 g of slow-release fertilizer pellets containing 15 mg (nitrate and ammoniacal nitrogen), 3.9 mg of phosphorous pentoxide, 9.1 mg of potassium oxide, 1.5 mg of magnesium oxide (plus microelements) every two months into rockpools containing Sabellaria alveolata. The temporal variability in nutrient enrichment had no effect on the abundance on Sabellaria alveolata during the 20-month study period.

Eutrophication may support the growth of green algae such as Ulva spp. Dubois et al. (2006) reported that algal epibionts reduced recruitment of Sabellaria alveolata, with potential but unknown impacts on long-term maintenance of reefs.

Sensitivity assessment. The evidence from direct nutrient enrichment (Bertocci et al., 2017) suggests that Sabellaria alveolata is not sensitive to this pressure. However, the possible but uncertain effects of green algal overgrowth of the reef (Dubois et al., 2006) suggest that eutrophication may adversely affect the reef. However, there is 'Insufficient evidence' on which to base an assessment at present.  

Organic enrichment

Benchmark. A deposit of 100 gC/m²/yr (Organic enrichment pressure definition).

Evidence

No 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 alveolata would be able to consume re-suspended particulate organic matter.  This conclusion is supported by the enhanced growth rates observed in the congener Sabellaria spinulosa 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).  Hence, sensitivity is recorded as 'Not sensitive'.

Physical loss (to land or freshwater habitat)

Benchmark. A permanent loss of existing saline habitat within the site (Physical loss pressure definition). 

Evidence

All marine habitats and benthic species are considered to have ‘None' resistance to this pressure and to be unable to recover (resilience is 'Very Low') 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 and the rate and confidence in resilience are not assessed.

Physical change (to another seabed type)

Benchmark. Permanent change from sedimentary or soft rock substrata to hard rock or artificial substrata, or vice versa (Physical change in subtratum type pressure definition).

Evidence

Sabellaria alveolata generally requires hard substrata on which to form reefs, but these must be in areas with a good supply of suspended coarse sediment. Reefs therefore commonly form on areas of rock or boulders surrounded by sand: the basis for this biotope group. Sabellaria alveolata reefs may also form on stable sediments.  Larsonneur et al., (1984), working in the Bay of St Michel in Normandy, noted that the sand mason Lanice conchilega can stabilise sand well enough to allow subsequent colonisation by Sabellaria alveolata. Settlement is also enhanced by the presence of existing colonies or their dead remains (Holt et al., 1998). Colonies on sand or other sediment may, however, be more short-lived as sediment mobility will disrupt reef formation (Holt et al., 1998).

Sensitivity assessment. The biotope classification refers specifically to rock habitats and therefore a change in sediment type from a rock substratum to a sedimentary substratum would significantly alter the biotope type. Due to the reduction in habitat suitability for reefs following permanent or prolonged substratum type changes, Sabellaria alveolata are judged to have ‘Low’ resistance to this pressure, resilience is Very low (the pressure is a permanent change) and sensitivity is assessed as High.

Physical change (to another sediment type)

Benchmark. Permanent change in one Folk class (based on UK SeaMap simplified classification) (Physical change in sediment type pressure definition). 

Evidence

Sabellaria alveolata generally requires hard substrata on which to form reefs, but these must be in areas with a good supply of suspended coarse sediment. Reefs therefore commonly form on areas of rock or boulders surrounded by sand: the basis for this biotope group. Sabellaria alveolata reefs may also form on stable sediments.  Larsonneur et al. (1984), working in the Bay of St Michel in Normandy, noted that the sand mason Lanice conchilega can stabilise sand well enough to allow subsequent colonisation by Sabellaria alveolata. Settlement is also enhanced by the presence of existing colonies or their dead remains (Holt et al. 1998).

There is some evidence that newly constructed groynes off Morecambe have resulted in a coarser sediment regime which has allowed Sabellaria alveolata to colonise boulder and cobble grounds in place of Mytilus edulis, which was previously dominant (Lumb, pers. comm.; Andrews, pers. comm., cited from Holt et al., 1998). Increases in coarse fractions are therefore considered to be beneficial for this species, particularly where these are stable. However, an increase in finer sediment has the potential to restrict development and high levels of silt in sediments will not provide a suitable substratum for colonization.

Sensitivity assessment. As this biotope specifically occurs on hard substratum, the pressure benchmark (a change in sediment type) is considered Not Relevant to the biotope group. 

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) (Removal of substratum pressure definition). 

Evidence

The removal of substratum down to 30 cm depth is likely to remove the whole Sabellaria alveolata reef within the extraction footprint.  At an expert workshop convened to assess the sensitivity of marine features to support MCZ planning, Sabellaria alveolata reefs were assessed as having no resistance to extraction of the feature (benchmark was the removal of feature/substrate to 50 cm depth) (Tillin et al., 2010).

Sensitivity assessment. As Sabellaria alveolata reefs are surface features they will be directly removed by extraction of the reef to 30 cm depth.  Resistance to this pressure is therefore assessed as ‘None’.  Resilience 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. Confidence in this assessment is assessed as 'High' due to the incontrovertible nature of the pressure. 

Abrasion / disturbance of the surface of the substratum or seabed

Benchmark. Damage to surface features (e.g. species and physical structures within the habitat) (Surface abrasion/disturbance pressure definition).

Evidence

Vorberg (2000) used video cameras to study the effect of shrimp fisheries on Sabellaria alveolata reefs to examine damage from fishing activities in the Wadden Sea. The imagery showed that the 3 m 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 (4.4 mm after removal of 2 cm of surface) than undisturbed growth (0.7 mm) and indicated that, as long as the reef is not completely destroyed, recovery can occur rapidly. These recovery rates are as a result of short-term effects following once-only disturbance. 

Cunningham et al. (1984) examined the effects of trampling on Sabellaria alveolata reefs. The reef recovered within 23 days from the effects of trampling (i.e. treading, walking or stamping on the reef structures) repairing minor damage to the worm tube porches. However, severe damage, estimated by kicking and jumping on the reef structure, resulted in large cracks between the tubes, and removal of sections (ca 15x15x10 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. At another site, a smaller section (10x10x10 cm) was lost, but after 23 days the space was already smaller due to rapid growth. Plicanti et al. (2016) further investigated the impact of trampling of on Sabellaria alveolata reefs in Portugal. By walking over sections of the reef once, twice or three times (low, medium and high intensity trampling), they demonstrated that even low intensity disturbance caused significant damage to the reef by reducing the amount of intact tubes. In contrast to previous studies, they demonstrated that, despite some increases, the percentage cover of intact tubes had not recovered to control levels two months after the disturbance and remained significantly reduced at sites exposed to medium and high intensity trampling.

Sensitivity assessment. For some impacts such as trampling and abrasion that leave behind large proportions of intact reef, recovery from a single event can occur within two years by rapid recolonization and expansion into damaged areas, facilitated by adults. Therefore, resistance is assessed as ‘Medium’, resilience is assessed as ‘High’ and sensitivity as ‘Low’. The scale and intensity of impacts would influence the level of resistance and the mechanism of recovery. Where reefs suffer extensive spatial damage requiring larval settlement to return to pre-impact conditions then recovery would be prolonged (years).

Penetration or disturbance of the substratum subsurface

Benchmark. Damage to sub-surface features (e.g. species and physical structures within the habitat) (Sub-surface penetration pressure definition).

Evidence

This pressure will result in the surface disturbance effects outlined above but effects will be compounded by the penetration and sub-surface damage aspect of this pressure. No empirical evidence was found to assess impacts however it is considered that the deeper and more significant the damage, the higher the risk of removing complete tubes and limiting recovery of the reefs.

Sensitivity assessment. Based on the evidence cited above for abrasion, resistance was assessed as ’Low’ (taking into account deeper penetration of the disturbance), recovery was assessed as ‘Medium’ (2-10 years) to take into account that larval recruitment may be necessary for the reef structure to recover although small, localised areas of repair would take place within months.  Sensitivity is, therefore assessed as ‘Medium’. 

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 (Suspended sediment pressure definition).

Evidence

Sabellaria alveolata do not rely on light penetration for photosynthesis and their visual perception is believed to be limited.  Changes in light penetration or attenuation associated with this pressure are, therefore, not relevant to the Sabellaria alveolata reef biotope, however alterations in the availability of suspended organic matter that can be used as food and the availability of suspended sediment for tube building could either increase or decrease habitat suitability for Sabellaria alveolata reefs.

The effect of increased seston concentration on Sabellaria alveolata clearance rates was investigated by Dubois et al. (2009). The range of experimental suspended particulate matter (SPM) concentrations (65-153.8 mg/l) corresponds to clear to medium turbidity at the pressure benchmark scale. The number of polychaetes actively feeding increased between  SPM 6.5-12.3 mg/l and no change was observed between SPM 12.3 and 55.5 mg/l.  At higher levels of SPM clearance rates were reduced, the decline in filter feeding efficiency (measured as a clearance rate) declined at around SPM 45 mg/l and thereafter remained relatively stable.

Tillin (2010) used logistic regression to develop statistical models that indicate how the probability of occurrence of 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.  The upper levels of these modelled optima broadly correspond with observations by Cayocca et al. (2008, cited in Dubois et al. 2009) who recorded SPM peaks ranging between 200 and 1000 mg/l depending on the flow and ebb conditions, in the vicinity of the largest Sabellaria alveolata reef in the Bay of Mont-Saint-Michel. Outside of these peaks, the SPM remained around 50 mg/l the level at which Dubois et al. (2009) recorded changes in clearance rate.

Sensitivity assessment. Sabellaria alveolata is adapted to turbid systems and can maintain its filtering activity under high seston loads (Dubois et al., 2009).   A supply of suspended sediment is a requirement for the development of reefs (Cunningham et al. 1984). Based on Cayocca et al. (2008, cited in Dubois et al., 2009) the normal range of SPM in which Sabellaria alveolata reefs occur is probably in the intermediate range (based on UKTAG, 2014 ranks) with increases in SPM at the beginning of ebb and flow periods.  It is therefore considered that Sabellaria alveolata reef biotopes are ‘Not sensitive’ to increases in peak suspended sediment concentration to the medium turbidity level (100-300 mg/l)  at the pressure benchmark. However, if the increase was constant then reductions in filtration efficiency may negatively affect a proportion of the population, resistance was therefore assessed as ‘Medium’ and recovery as ‘High’ following habitat recovery. Sensitivity is therefore considered to be ‘Low’.  A reduction from intermediate levels to clear (<10 mg/l) where the reduction is due to a reduced supply of organic matter and particulate matter suitable for tube building and food may restrict reef development and reduce the food supply to this species. Resistance was assessed as ‘Low’ and recovery as ‘Medium’ so that sensitivity is considered to be ‘Medium’. 

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 (Smothering pressure definition).

Evidence

Sabellaria alveolata do not rely on light penetration for photosynthesis and their visual perception is believed to be limited.  Changes in light penetration or attenuation associated with this pressure are, therefore, not relevant to the Sabellaria alveolata reef biotope, however alterations in the availability of suspended organic matter that can be used as food and the availability of suspended sediment for tube building could either increase or decrease habitat suitability for Sabellaria alveolata reefs.

The effect of increased seston concentration on Sabellaria alveolata clearance rates was investigated by Dubois et al. (2009). The range of experimental suspended particulate matter (SPM) concentrations (65-153.8 mg/l) corresponds to clear to medium turbidity at the pressure benchmark scale. The number of polychaetes actively feeding increased between  SPM 6.5-12.3 mg/l and no change was observed between SPM 12.3 and 55.5 mg/l.  At higher levels of SPM clearance rates were reduced, the decline in filter feeding efficiency (measured as a clearance rate) declined at around SPM 45 mg/l and thereafter remained relatively stable.

Tillin (2010) used logistic regression to develop statistical models that indicate how the probability of occurrence of 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.  The upper levels of these modelled optima broadly correspond with observations by Cayocca et al. (2008, cited in Dubois et al. 2009) who recorded SPM peaks ranging between 200 and 1000 mg/l depending on the flow and ebb conditions, in the vicinity of the largest Sabellaria alveolata reef in the Bay of Mont-Saint-Michel. Outside of these peaks, the SPM remained around 50 mg/l the level at which Dubois et al. (2009) recorded changes in clearance rate.

Sensitivity assessment. Sabellaria alveolata is adapted to turbid systems and can maintain its filtering activity under high seston loads (Dubois et al., 2009).   A supply of suspended sediment is a requirement for the development of reefs (Cunningham et al. 1984). Based on Cayocca et al. (2008, cited in Dubois et al., 2009) the normal range of SPM in which Sabellaria alveolata reefs occur is probably in the intermediate range (based on UKTAG, 2014 ranks) with increases in SPM at the beginning of ebb and flow periods.  It is therefore considered that Sabellaria alveolata reef biotopes are ‘Not sensitive’ to increases in peak suspended sediment concentration to the medium turbidity level (100-300 mg/l)  at the pressure benchmark. However, if the increase was constant then reductions in filtration efficiency may negatively affect a proportion of the population, resistance was therefore assessed as ‘Medium’ and recovery as ‘High’ following habitat recovery. Sensitivity is therefore considered to be ‘Low’.  A reduction from intermediate levels to clear (<10 mg/l) where the reduction is due to a reduced supply of organic matter and particulate matter suitable for tube building and food may restrict reef development and reduce the food supply to this species. Resistance was assessed as ‘Low’ and recovery as ‘Medium’ so that sensitivity is considered to be ‘Medium’. 

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 (Smothering pressure definition).

Evidence

Sabellaria alveolata was reported to survive short-term burial for days and even weeks in the south west of England as a result of storms that altered sand levels up to two meters, they were, however, killed by longer-term burial (Earll & Erwin 1983). Sabellaria alveolata has been identified as sensitive to changes in sediment regime in the Mediterranean Gulf of Valencia, Spain, where Sabellaria alveolata populations were lost as a result of sand level rise resulting from the construction of seawalls, marinas/harbours, and beach nourishment projects (Porras et al., 1996). It is likely that the length of survival, while dependent on the length of burial, may be influenced by temperatures and oxygen levels so that seasonality and the depth and character of overburden partially determine sensitivity.

Sensitivity assessment.  Natural events such as storms may lead to episodic burial by coarse sediments with subsequent removal by water action and the degree of mortality will depend on a number of factors including the length of burial. As fine sediments may be relatively cohesive and as water and air penetration are limited, the addition of an overburden of 30 cm is considered to potentially lead to some mortality if large areas are impacted. Resistance is therefore assessed as ‘Low’ and recovery is assessed as ‘Medium’, and sensitivity to this pressure is categorised as ‘Medium’.

Litter

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

Evidence

Not assessed.

Electromagnetic changes

Benchmark. A local electric field of 1 V/m or a local magnetic field of 10 µT (Electromagnetic pressure definition).

Evidence

Evidence on the effect of electromagnetic fields (EMFs) on benthic organisms is still severely lacking. Some studies have investigated the effect of anthropogenically induced EMFs on benthic invertebrates at intensities ranging between 2 nT and 40 mT, which is often much higher than in-situ measurements from subsea cables. While some report changes to behaviour, physiology, reproduction, development, immunology, cytotoxicity and orientation, others demonstrate no effect from exposure to the EMF (Albert et al., 2020; Hutchison et al., 2020), depending on the study species and duration and intensity of exposure. There have been no studies investigating the effect of EMFs at the population or community level for benthic organisms. 

No studies have examined the effect of EMFs on Sabellaria alveolata. However, one study was performed on another reef forming annelid, Ficopomatus enigmaticus (Oliva et al., 2023). Sperm cells from this species were exposed to 0.5 and 1.0 mT of static magnetic field. After only three hours of exposure, sperm fertilization rate was reduced and significant increases in DNA damage and mitochondrial activity indicative of a stress response were reported. However, there is 'Insufficient evidence' on which to base an assessment of the likely sensitivity of Sabellaria reefs to EMFs.

Underwater noise changes

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

Evidence

No evidence.

Introduction of light or shading

Benchmark. A change in incident light via anthropogenic means (Introduced light or shade pressure definition).

Evidence

Since 2016, research on artificial light at night (ALAN) has expanded considerably in the marine and coastal environment. Light was previously assumed to be of low ecological significance in subtidal and intertidal habitats, but there is now evidence that ALAN is widespread in the marine environment, with biologically relevant levels of light penetrating to depths of up to 50m (Davies et al., 2020; Smyth et al., 2021). ALAN can alter biological processes across taxa and at multiple levels of organisation. Documented responses include disruption of diel and circalunar rhythms, changes in activity and foraging, altered predator–prey interactions, shifts in community composition, and impacts on algal growth and phenology (Davies et al., 2014, 2015; Gaston et al., 2017; Tidau et al., 2021; Lynn et al., 2022; Marangoni et al., 2022; Miller et al., 2023; Ferretti et al., 2025). Evidence for benthic habitats and assemblages specifically is beginning to emerge (e.g. Trethewy et al., 2023; Schaefer et al., 2025), but remains limited and fragmented, often focusing on single taxa or short-term experiments. Mortality thresholds, long-term consequences, and responses at the biotope scale are rarely addressed, and there are major gaps around indirect effects such as trophic cascades or habitat modification.

Sensitivity assessment. Given the rapid expansion of the evidence base but the continuing lack of data at the level of individual biotopes, resistance and resilience cannot be robustly assessed. Sensitivity is therefore recorded as 'Insufficient evidence'.

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 (Barrier to species movement pressure definition).

Evidence

Barriers that reduce the degree of tidal excursion may reduce the supply of Sabellaria alveolata larvae moving landwards to suitable habitats from source populations. However, the presence of barriers may enhance local population supply by preventing the seaward loss of larvae. The residual tidal currents in Bay of Mont-Saint-Saint Michel (France) naturally prevent the loss of larvae from the bay and are believed to enhance settlement locally (Dubois et al., 2007). 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’.

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 (Death for collision pressure definition).

Evidence

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

Visual disturbance

Benchmark. The daily duration of transient visual cues exceeds 10% of the period of site occupancy by the feature (Visual disturbance pressure definition). 

Evidence

Not relevant.

Genetic modification & translocation of indigenous species

Benchmark. Translocation of indigenous species or the introduction of genetically modified or genetically different populations of indigenous species may result in changes in the genetic structure of local populations, hybridization, or a change in community structure (Translocation pressure definition).

Evidence

Sabellaria alveolata is not farmed or translocated, therefore this pressure is 'Not relevant'.

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) (pathogen or disease pressure definition).

Evidence

No evidence found for pathogens or diseases impacting Sabellaria alveolata.

Removal of target species

Benchmark. Removal of species targeted by fishery, shellfishery or harvesting at a commercial or recreational scale (targeted removal pressure definition).

Evidence

Sabellaria alveolata biotopes may be removed or damaged by people harvesting species, such as mussels, within the biotope or by contact with 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 extraction of Sabellaria alveolata by bait digging is a possibility. Damage to colonies by people opening tubes with knives and removing the worms for use as fishing bait has been observed, though nowhere has this been seen on any intensive scale (references in Holt et al., 1998) in the UK. Only one study investigated the effect of direct harvesting on the recovery of Sabellaria alveolata reefs in Italy, where removal for bait has been observed. Storari et al. (2024) removed sections of the reef by taking various sized core samples (low, medium and high intensity disturbance). After 20 months, the reef sites from which the low and high intensity harvesting occurred recovered to the same patch size as the unharvested site. The site from which the medium sized core was taken demonstrated larger Sabellaria alveolata patches compared to all others, increasing by 36%. This could suggest that Sabellaria alveolata benefits from intermediate levels of disturbance, perhaps through decreased localised intra-specific competition facilitating the bioconstruction from surviving adults, which aligns with findings from Cunningham et al. (1984) and Vorberg (2000).

No evidence was found for trophic or other ecological interactions between commercially targeted species and Sabellaria alveolata.

Sensitivity assessment. Sabellaria alveolata is not commercially targeted in the UK. However, hand gathering may occur at a low intensity, and larger scale harvesting such as core-taking, scraping, or shovelling can remove entire portions of the reef. Therefore, resistance is assessed as ‘Medium’. As full recovery was documented 20 months after harvesting, recovery is assessed as ‘High’. Sensitivity of this biotope is therefore considered ‘Low’.  

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 (non-targeted removed pressure definition).

Evidence

Sabellaria alveolata 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.  Sabellaria alveolata creates the biogenic reefs that characterise this biotope, removal of this species as by-catch would, therefore, remove the biotope.  No evidence was found for key trophic or other ecological interactions between other species within the biotope and Sabellaria alveolata.

Sensitivity assessment. Removal of the worms and tubes as by-catch would remove the biotope and hence this group is considered to have ‘None’ resistance to this pressure and to have ‘Medium’ recovery. Sensitivity is, therefore ‘Medium’. 

The American slipper limpet, Crepidula fornicata

Evidence

The slipper limpet Crepidula fornicata was recorded on intertidal Sabellaria alveolata reefs in Champeaux, west Cotentin coast, northern France, at a low density of ca 0.75+/-2 /m² (Schlund et al., 2016). Powell-Jennings & Calloway (2018) reported that Crepidula had a preference for hard grounds colonized by Sabellaria alveolata in Swansea Bay, south Wales, with over 80% of the records of Crepidula associated with the Sabellaria alveolata reef. However, no evidence of their relationship was available, and they may be in competition or facilitate each other's presence (Powell-Jennings & Calloway, 2018). 

Sensitivity assessment. The evidence above suggests that Crepidula fornicata has the potential to colonize intertidal Sabellaria alveolata reefs. There is no evidence to suggest that Crepidula has a detrimental effect on the reefs. The presence of sand combined with wave exposure and storm-related scour may limit the ability of Crepidula to colonize the habitat and its density. Therefore, the resistance to colonization by Crepidula fornicata is assessed as ‘Medium’. Resilience is likely to be 'Very low' as a bed of Crepidula fornicata would need to be removed (by human intervention) before recovery could begin. Therefore, sensitivity is considered to be ‘Medium’. However, the confidence in this assessment is 'Low' due to the lack of evidence of deleterious effects. 

The carpet sea squirt, Didemnum vexillum

Evidence

The carpet sea squirt Didemnum vexillum (syn. Didemnum vestitum; Didemnum vestum) is a colonial ascidian with rapidly expanding populations that have invaded most temperate coastal regions around the world (Kleeman, 2009; Stefaniak et al., 2012; Tillin et al., 2020). It is an ‘ecosystem engineer’ that can change or modify invaded habitats and alter biodiversity (Griffith et al., 2009; Mercer et al., 2009). Didemnum vexillum has colonized and established populations in the northeast Pacific, Canadian and USA coast; New Zealand; France, Spain, and the Wadden Sea, Netherlands; the Mediterranean Sea and Adriatic Sea (Bullard et al., 2007; Coutts & Forrest, 2007; Dijkstra et al., 2007; Valentine et al., 2007a; Valentine et al., 2007b; Lambert, 2009; Hitchin, 2012; Tagliapietra et al., 2012; Gittenberger et al., 2015; Vercaemer et al., 2015; Mckenzie et al., 2017; Cinar & Ozgul, 2023; Holt, 2024). In the UK, Didemnum vexillum has colonized Holyhead marina and Milford Haven, Wales; the west coast of Scotland (marinas around Largs, Clyde, Loch Creran and Loch Fyne), South Devon (Plymouth, Yealm, and Dartmouth estuaries), the Solent, northern Kent, Essex, and Suffolk coasts (Griffith et al., 2009; Lambert, 2009; Hitchin, 2012; Michin & Nunn, 2013; Bishop et al., 2015; Mckenzie et al., 2017; Tillin et al., 2020, Holt, 2024; NBN, 2024).

Although a widespread invader, Didemnum vexillum has a limited ability for natural dispersal since the pelagic larvae remain in the water column for a short time (up to 36 hours). Therefore, it has a short dispersal phase that can allow the species to build localized populations (Herborg et al., 2009; Vercaemer et al., 2015; Holt, 2024). However, Bullard et al. (2007) suggested that Didemnum vexillum can form new colonies asexually by fragmentation. Colonies can produce long tendrils from an encrusting colony, which can fragment, disperse and settle, attaching to suitable hard substrata elsewhere (Bullard et al., 2007; Lambert, 2009; Stefaniak & Whitlatch, 2014). A fragmented colony can spread naturally for up to three weeks transported by ocean currents, attached to floating seaweed, seagrass or other floating biota, or as free-floating spherical colonies (Bullard et al., 2007; Lengyel et al., 2009; Stefaniak & Whitlatch, 2014; Holt, 2024). Fragments can reattach to suitable substrata within six hours of contact. Fragments have the potential to disperse around 20 km before reattachment (Lengyel et al., 2009). Valentine et al. (2007a) reported that colonies of Didemnum vexillum enlarged by 6 to 11 times by asexual budding after 15 days and enlarged 11 to 19 times after 30 days. Valentine et al. (2007a) concluded fragments could successfully grow, survive, and help to spread Didemnum vexillum.

While natural fragmentation of tendrils is thought to allow Didemnum vexillum to invade longer distances and increase its dispersal potential, Stefaniak & Whitlatch (2014) found that only one tendril out of 80 reattached to the flat, bare substrata used in their study, because tendrils required an extensive (at least eight hour) period of contact to reattach. Stefaniak & Whitlatch (2014) suggested that once fragmented from a colony, the success of tendril reattachment was limited, and reattachment was not a major contributor to the invasive success of Didemnum vexillum. However, Stefaniak & Whitlatch (2014) also found that larvae-packed tendril fragments may increase natural dispersal distance, reproduction, and invasive success of Didemnum vexillum, and increase the distance larvae can travel. Not all colonies produce tendrils at all locations.

Human-meditated transport via aquaculture facilities, boat hulls, commercial fishing vessels, and ballast water is probably the most important vector that has aided the long-distance dispersal of Didemnum vexillum and explains its prevalence in harbours and marinas (Bullard et al., 2007; Dijkstra et al., 2007; Griffith et al., 2009; Herborg et al., 2009). Fragmentation of colonies during transport or human disturbance (such as trawling or dredging) could indirectly disperse the species and enable it to find suitable conditions for establishment (Herborg et al., 2009). For example, in oyster farms in British Columbia, large fragments of Didemnum sp. come off oyster strings when they are pulled out of water and other fragments can be pulled off oysters and mussels and thrown back into the water, which is likely to aid dispersal of the invasive species (Bullard et al., 2007). Dijkstra et al. (2007) hypothesised that Didemnum sp. was introduced to the Gulf of Maine with oyster aquaculture in the Damariscotta River and transported via Pacific oysters.

Didemnum vexillum was likely introduced into the UK from northern Europe or Ireland via poorly maintained or not antifouled vessels, movement of contaminated shellfish stock and aquaculture equipment, or via marine industries such as oil, gas, renewables, and dredging (Holt, 2024). Recent evidence from genetic material suggests that human-mediated dispersal, between marinas and shellfish culture sites, is the most likely pathway for connectivity of Didemnum vexillum populations throughout Ireland and Britain (Prentice et al., 2021; Holt, 2024). Didemnum vexillum can disperse away from artificial substrata, invading and colonizing natural substrata in surrounding areas (Tillin et al., 2020). Holt (2024) noted that Didemnum vexillum had not spread as far as feared in the UK since it was first recorded. The current evidence of Didemnum vexillum’s ability to spread on natural habitats in this area is sparse and often conflicting, complicated by genetics and its apparent variable habitat preferences and tolerances and its variable ability to adapt to ‘new’ conditions (Holt 2024).

Didemnum vexillum has a seasonal growth cycle that is influenced by temperature (Valentine et al., 2007a). In warmer months (June and July) colonies may be large and well-developed encrusting mats. Populations experience more rapid growth from July to September sometimes continuing into December. Colonies begin to decline in health and ‘die-off’ when temperatures drop below 5°C during winter months from around October to April (Gittenberger, 2007; Valentine et al., 2007a; Herborg et al., 2009). Cold water months cause colonies to regress and reduce in size, yet they often regenerate as temperatures warm (Griffith et al., 2009; Kleeman, 2009, Mercer et al., 2009), although some populations may not survive winter at all (Dijkstra et al., 2007). The early growth phase, from May to July, is initiated by smaller colonies developing from remnants of colonies that survived the cold water (Valentine et al., 2007a). The seasonal growth cycle is also likely influenced by location. For example, the Didemnum sp. growth cycle for colonies in Sandwich tide pool (temperature range from -1 °C to 24 °C, with daily fluctuations), probably does not occur in deep offshore subtidal habitats in Georges Bank (annual temperature range from 4 °C to 15°C, and daily fluctuations are minimal) (Valentine et al., 2007a). Larval release and recruitment typically occur between 14 to 20°C and slow or cease below 9 to 11°C as summer ends (Griffith et al., 2009; Mckenzie et al., 2017). In New Zealand, recruitment occurs from November to July, where the highest average temperatures were recorded in February (18 to 22°C) and the lowest average temperatures were recorded in July (9 to 10°C) (Fletcher et al., 2013a). In this New Zealand study, higher water temperatures were associated with a higher level of recruitment (Fletcher et al., 2013a).

Didemnum vexillum requires suitable hard substrata for successful settlement and the establishment of colonies. It can grow quickly and establish large colonies of dense encrusting mats on a variety of hard substrata (Valentine et al., 2007a; Griffith et al., 2009; Lambert, 2009; Groner et al., 2011; Cinar & Ozgul, 2023). Gittenberger (2007) stated that invasive Didemnum sp. was a threat to native ecosystems because of its ability to overgrow virtually all hard substrata present. Suitable hard substrata can include rocky substrata such as bedrock gravel, pebble, cobble, or boulders or artificial substrata such as a variety of maritime structures such as pontoons, docks, wood and metal pilings, chains, ropes and moorings, plastic and ship hulls and at aquaculture facilities (Valentine et al., 2007 a&b; Bullard et al., 2007; Griffith et al., 2009; Lambert, 2009; Tagliapietra et al., 2012; Tillin et al., 2020). Didemnum vexillum has been reported colonizing these types of hard substrata in the USA, Canada, northern Kent, and the Solent (Bullard et al., 2007; Valentine et al., 2007a; Valentine et al., 2007b; Hitchin, 2012; Vercaemer et al., 2015; Tillin et al., 2020).

Didemnum vexillum has the ability to rapidly overgrow and displace on other sessile organisms such as other colonial ascidians (Ciona intestinalis, Styela clava, Ascidiella aspera, Botrylloides violaceus, Botryllus schlosseri, Diplosoma listerianium and Aplidium spp.), bryozoan, hydroids, sponges (Clione celata and Halichrondria sp.), anemone (Diadumene cincta), calcareous tube worms, eelgrass (Zostera marina), kelp (Laminaria spp. and Agarum sp.), green algae (Codium fragile subsp. fragile), red algae (Plocamium, Chondrus crispus and bush weed Agardhiella subulata), brown algae (Ascophyllum nodosum, Sargassum, Halidrys, Fucus evanescens and Fucus serratus), calcareous algae (Corallina officinalis), mussels (Mytilus galloprovincialis, Perna canaliculus  and Mytilus edulis), barnacles, oysters (Magallana gigas, Ostrea edulis and Crassostrea virginica), sea scallops (Placopecten magellanicus), or dead shells (Dijkstra et al., 2007; Gittenberger, 2007; Valentine et al., 2007a; Valentine et al., 2007b; Griffith et al., 2009; Carman & Grunden, 2010; Dijkstra & Nolan, 2011; Groner et al., 2011; Hitchin, 2012; Tagliapietra et al., 2012; Minchin & Nunn, 2013; Gittenberger et al., 2015; Long & Groholz, 2015; Vercaemer et al., 2015).

In contrast, Didemnum vexillum’s preference for sheltered conditions, established colonies observed in Georges Bank and Long Island Sound were exposed to moderately strong tidal currents (1 to 2 knots; ca 0.5 to 1 m/s recorded at both sites) that may mobilise sediment (Valentine et al., 2007b; Mercer et al., 2009; Tillin et al., 2020). However, Valentine et al. (2007b) describe the substratum as immobile, presumably consolidated gravel, cobbles, and pebbles. Kleeman (2009) stated that the presence of a consistent mild wave action or ‘swash zone’ appears to favour Didemnum sp. establishment in the intertidal. Although some evidence suggests that waves and currents can facilitate the fragmentation and spread of Didemnum vexillum (Mckenzie et al., 2017), the tidal current velocities at some sites where Didemnum vexillum has been reported (for example, New England, where current velocities reach up to around 3 m/s) is lower than the current velocity required for the dislodgement of Didemnum vexillum fragments (around 7.6 m/s) (Reinhardt et al., 2012). This suggests that not all tidal currents are likely to dislodge Didemnum vexillum fragments. When on boat hulls the species can experience higher current velocities which is enough to cause dislodgement (Reinhardt et al., 2012).  

Sensitivity assessment. Didemnum vexillum has not been recorded colonizing Sabellaria reefs or associating with Sabellaria alveolata. The presence of rocky substrata in this Sabellaria biotope could provide a suitable hard surface for the successful colonization of Didemnum vexillum. Furthermore, Didemnum vexillum has been recorded in moderately strong currents (Valentine et al., 2007b; Mercer et al., 2009; Tillin et al., 2020) and is predicted to survive stronger currents, as the current velocity which will dislodge Didemnum vexillum fragments is around 7.6 m/s (Reinhardt et al., 2012). Therefore, Didemnum vexillum could colonize areas in which intertidal Sabellaria reefs inhabit as their optimum tidal current was modelled at less than 3 m/s (Tillin, 2010).

Holt (2024) noted that Didemnum vexillum had not spread as far as feared in the UK since it was first recorded, and no evidence has been found on the potential effects of Didemnum sp. on Sabellaria alveolata. If Didemnum sp. could gain a 'foothold' it might overgrow, smother or cause mortality on the Sabellaria reefs. However, despite reports of other biogenic reefs, such as mussel and oyster beds, being suitable substrata, Tillin et al. (2010) considered intertidal Sabellaria alveolata reefs to be too well drained for Didemnum vexillum to survive. Therefore, a resistance of 'Medium' (some, <25% mortality) is suggested as a precaution in case Didemnum vexillum can colonize the biotope, but with 'Low' confidence due to the lack of direct evidence. Resilience is assessed as 'Very low' as recovery would require the physical removal of Didemnum sp., so sensitivity is assessed as 'Medium'. 

The Pacific oyster, Magallana gigas

Evidence

The Pacific oyster, Magallana (syn. Crassostrea) gigas, is native to warm temperate regions from the north-west Pacific to Japan and north-east Asia, including Cape Mariya (Russia) to Hong Kong (China) (Carrasco & Baron, 2010; GBNNSS, 2011, 2012). It is a fast-growing and tolerant species that has become a successful invader in the coastal waters of all continents, aside from Antarctica (Wrange et al., 2010; Carrasco & Baron, 2010; Padilla, 2010). Magallana gigas is recognised as a beneficial and important species in aquaculture worldwide (Padilla, 2010). It was initially introduced for aquaculture in Europe and the UK in the 1960s due to a decline in the Portuguese oyster (Crassostrea angulata) and the European flat oyster (Ostrea edulis) (Spencer et al., 1994; GBNNSS, 2011, 2012; Humphreys et al., 2014 cited in Alves et al., 2021; Hansen et al., 2023).

Since introduction, the species has invaded and established self-sustaining natural populations throughout Europe from the North Sea, Wadden Sea and Scandinavian coastlines to the Atlantic coastlines of Spain and Portugal, as well as the Mediterranean and Adriatic Sea (Wrange et al., 2010; GBNNSS, 2011, 2012; Ezgeta-Balic et al., 2019; Spagnolo et al., 2019; Bergstrom et al., 2021; Hansen et al., 2023). In the UK, the species predominantly occurs around the southern and western coastlines (OBIS, 2024; NBN, 2024). Shipping activity has also been associated with the introduction of Magallana gigas in the north-eastern Adriatic Sea, where it was not introduced for aquaculture (Ezgeta-Balic et al., 2019). It was also suggested that some Magallana gigas populations were established in south-west England from France possibly via fouling on ships (GBNNSS, 2011, 2012; Padilla, 2010; Ezgeta-Balic et al., 2019).

Magallana gigas has a high fecundity, a long-lived pelagic larval phase (2 to 4 weeks) and can produce up to 200 million eggs during spawning (Herbert et al., 2012, 2016; Alves et al., 2021; Wood et al., 2021; Hansen et al., 2023). Hence, as a broadcast spawner, it has a high dispersal potential of more than 1000 km (Padilla, 2010; Wood et al., 2021). Larval mortality can be as large as 99%, as larvae are sensitive to environmental conditions (Alves et al., 2021). However, adults are long-lived so populations can survive with infrequent recruitment (Padilla, 2010). Larval dispersal and mass spawning events have facilitated the settlement and establishment of Pacific oysters, as seen in the Oosterschelde estuary, Netherlands (Hansen et al., 2023). It has been suggested that the spread of the Pacific oyster in Scandinavia is due to northward larval drift on tidal and wind-driven currents (Hansen et al., 2023). Wood et al. (2021) suggested that larval dispersal of the Pacific oyster from populations within and outside the UK was possible via unaided (passive) transport by currents, but that aquaculture and offshore structures (e.g. windfarms) increased the risk of the invasive species spreading and the geographical extent of spread.

Magallana gigas is an ecosystem engineer and can dramatically change habitat structure when it invades. Once successfully settled, groups of Pacific oysters may form dense aggregations, potentially forming a reef, which in some regions can reach densities of 700 individuals m² (Herbert et al., 2012, 2016). Once, the density of live or dead Pacific oysters reaches or exceeds 200 ind./m² little of the underlying substratum remains visible (Herbert et al., 2016). These reefs can stabilize the sediment surface locally (Troost, 2010). When such reefs are formed or, particularly when the species colonizes soft sediments such as mud or sand, it can change and affect local communities, by creating hard substrata for mobile species, which might not otherwise be present before the invasion (Padilla, 2010). However, Hansen et al. (2023) suggested that where the Pacific oyster occurs sporadically, no immediate ecosystem risk is observed.

Magallana gigas also colonizes littoral intertidal biogenic reefs formed by the blue mussel Mytilus edulis or honeycomb worm Sabellaria alveolata (GBNNSS, 2011, 2012; Kochmann, 2012; Kochmann et al., 2013; Herbert et al., 2016; Tillin et al., 2020). The colonization and overgrowth of Magallana gigas may have impacts on Sabellaria alveolata and its habitat formation (Herbert et al., 2012, 2016). Pacific oysters may smother Sabellaria alveolata because it grows over tube ends and could outcompete it for space (Dubois et al., 2006; Desroy et al., 2011). Colonization by the Pacific oyster has been linked to the degradation and deterioration of Sabellaria reef health (Desroy et al., 2011). Desroy et al. (2011) reported several contributing factors, including an increase in silt deposits and fine particles in the sediment from pseudo-faeces produced by the oysters, which can cause increased sedimentation and nutrient enrichment (Green & Crowe, 2013). It has been suggested the increased sediment from oysters might explain why some species normally found in muddy-sand environments were present, further creating new species associations (Dubois et al., 2006).

Dubois et al. (2006) found that Magallana gigas had invaded some Sabellaria alveolata reefs in the Bay of Mont-Saint-Michel, France, resulting in densities of more than 100 oysters /m² on some of them. In this area, Sabellaria alveolata reefs were the only available hard substratum for settlement by the Pacific oyster. The study found that an intermediate covering of the Pacific oyster introduced greater species richness and heterogeneity of diversity on the Sabellaria reefs by creating hard substrata habitats and refuges for sessile or mobile species not usually present (Dubois et al., 2006). Green & Crowe (2013) found a lower percentage cover of Sabellaria alveolata on boulders colonized by Magallana gigas in Ireland. 

The presence of other filter feeders such as Magallana gigas increases trophic competition (Desroy et al., 2011, Green & Crowe, 2013). However, high densities of filter-feeding species alter the settlement of particulate matter and larvae as turbulence in the water column is increased (Green & Crowe, 2013). The physical structure of the oyster beds changes the hydrography and provides refuge from predators for oyster larvae, which increases their recruitment (Soniat et al., 2004). Pacific oysters might improve the recruitment of Sabellaria alveolata by increasing the probability of Sabellaria larvae swimming or sinking down the water column (Tillin et al., 2020). However, Dubois et al. (2006) reported that the abundance of smaller class sizes of Sabellaria alveolata was reduced in Sabellaria reefs with epibionts (Pacific oyster or Ulva spp.) indicating negative impacts of Pacific oysters on recruitment, although not as marked as in the presence of algae (Padilla, 2010; Tillin et al., 2020). Secondary impacts have also been reported, including increased recreational harvesting of the oysters on Sabellaria reefs, which led to reef trampling, physical damage and fragmentation (Dubois et al., 2006; Desroy et al., 2011). In the northern part of Bourgneuf Bay, France Magallana gigas was observed in rocky areas usually occupied by Sabellaria alveolata (Cognie et al., 2006; Herbert et al., 2012, 2016). Cognie et al. (2006) suggested that Magallana gigas could compete with Sabellaria alveolata for food and space, leading Herbert et al. (2012) to suggest that the Pacific oyster may prevent new colonization by Sabellaria alveolata.

Sensitivity assessment. The evidence above suggests Magallana gigas has the potential to colonize intertidal Sabellaria alveolata reefs. The presence of Magallana gigas at low densities can increase the species richness and diversity on Sabellaria alveolata reefs, but higher densities of the Pacific oyster may smother Sabellaria alveolata and outcompete it for space and food (Cognie et al., 2006; Dubois et al., 2006; Desroy et al., 2011). Magallana gigas may prevent the colonization of Sabellaria alveolata in some areas (Herbert et al., 2012). Therefore, the resistance to colonization by Magallana gigas is assessed as 'Low' as a precaution. Resilience is likely to be ‘Very low’ as the Magallana gigas population would need to be removed for recovery to occur. Therefore, the sensitivity of Sabellaria alveolata reef biotopes is assessed as ‘High’.

Wireweed, Sargassum muticum

Evidence

Wireweed, Sargassum muticum. Sargassum muticum is known to grow in the shallow subtidal around the UK, usually in areas sheltered from wave action. Therefore, while Sabellaria alveolata reefs may provide suitable attachment substrata, the exposed to moderately wave exposed nature in which this biotope is found may be unfavourable for Sargassum muticum. Its distribution is limited by the availability of hard substratum (e.g. stones >10 cm) and light (Staeher et al., 2000; Strong & Dring 2011; Engelen et al., 2015).  It is most abundant between 1 and 3 m below mean water, is a poor competitor under low light and only develops dense canopies in shallow areas (Engelen et al., 2015). Therefore, it is unlikely that Sargassum would colonize this biotope, hence it is 'Not sensitive'.

Wakame, Undaria pinnatifida

Evidence

Wakame, Undaria pinnatifida. Undaria pinnatifida is known to grow in the shallow subtidal around the UK but is usually found in areas sheltered from wave action, with a depth range of -1 to 4 m. Therefore, while Sabellaria alveolata reefs may provide suitable attachment substrata, the exposed to moderately wave exposed nature in which this biotope is found may be unfavourable for Undaria pinnatifida, hence it is 'Not sensitive'.

Other INIS

Evidence

Compass sea squirt, Asterocarpa humilis. While no evidence of colonization by Asterocarpa humilis has been reported, Sabellaria alveolata reefs were considered as potentially suitable habitat (albeit with low confidence; Tillin et al., 2020), based substratum suitably alone.

Red ripple bryozoan, Watersipora subatra. Sabellaria alveolata reefs were considered as potentially suitable habitats for Watersipora subatra as they provide suitable substrata on which it can attach. It is often found on the lower intertidal and shallow subtidal, and able to withstand various salinities and wave exposures (Tillin et al., 2020). However, no evidence of Watersipora subatra on Sabellaria alveolata reefs around the UK was found.

Red seaweed, Agarophyton vermiculophyllum. This invasive red seaweed shows associations with biogenic reefs, therefore, intertidal Sabellaria alveolata reefs within this biotope may be potentially suitable habitat (Till et al., 2020). However, no evidence of their occurrence on Sabellaria reefs has been documented. 

Orange striped anemone, Diadumene lineata. Tillin et al. (2020) suggested with low confidence that Sabellaria alveolata reefs may confer potentially suitable habitat for Diadumene lineata due to availability of attachment substrata and favourable environmental conditions. No evidence of Diadumene lineata colonizing Sabellaria reefs was found.

Sensitivity assessment. While this biotope may confer potentially suitable habitat for these species, there has been no direct evidence of their occurrence on Sabellaria alveolata reefs around the UK and Ireland. Therefore, there is ‘Insufficient evidence’ from which to assess the sensitivity of this biotope to these species.