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. Further detail.

Evidence

Caryophyllia smithii is a temperate cup coral that has a wide distribution from Norway to the Mediterranean and South Africa in the Atlantic, found in both shallow and deep waters, with records of this species between temperatures of 5 and 30°C. Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in seawater temperature. Therefore, unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns given that gamete release is most likely triggered by seasonal temperature increases (Tranter et al., 1982).

Alcyonium digitatum is a boreal species of octocoral recorded along the Atlantic Coasts of Europe from Portugal to Norway and Iceland, and along the North West Atlantic coasts, at sea surface temperatures between 5 and 20°C but mostly between 10 and 15°C (www.obis.org).  Across this latitudinal gradient species are likely to experience a range of temperatures from approx. 5 and 18°C (Sea temperature, 2015). 

Alcyonium digitatum spawns during the winter months (Hartnoll, 1975). Gametes are fertilized while in the water column and the embryos give rise to actively swimming lecithotrophic planulae that may have an extended pelagic life before they eventually settle (usually within one or two further days) and metamorphose to polyps (Matthews, 1917; Hartnoll, 1975). However, laboratory experiments have observed settlement failure to occur in unsuitable conditions (Hartnoll, 1975). As spawning occurs when sea temperatures are low there is the probability that spawning and settlement could be correlated to climatic conditions, therefore global warming could impact the reproduction and recruitment of Alcyonium digitatum. However, the combination of spawning in winter and the long pelagic lifespan may allow a considerable length of time for the planulae to disperse, settle and metamorphose ahead of the spring plankton bloom (Hartnoll, 1975).

The duration, dispersal and survival of planktonic larvae are dependent on several factors including temperature (O’Connor et al., 2007). O’Connor et al. (2007) reported planktonic larval duration to increase with temperature, therefore cold-water species could see an increase in planktonic larval duration under global warming trends. Larval survival has been reported to decrease exponentially with time (planktonic larval duration) (O’Connor et al., 2007). Elevated temperatures may increase the occurrence of octocoral diseases caused by pathogens that act opportunistically to attack hosts that are under stressful conditions. For example, Cerrano et al. (2000) reported that ecosystems in the Mediterranean are rapidly declining from extensive attacks by microorganisms correlating to elevated seawater temperatures. 

Echinus esculentus is a sea urchin distributed across the N.E. Atlantic from Iceland, north to Finmark, Norway and south to Portugal. Echinus esculentus is common on most coasts of the British Isles but absent from most of the east coast of England, the eastern English Channel and some parts of north Wales. Echinus esculentus has been recorded primarily between sea temperatures of 5 and 15°C (www.obis.org). 

Echinus esculentus occurred at temperatures between 0 and 18°C in Limfjord, Denmark (Ursin 1960). Temperature, photoperiod and food availability are considered to be factors that control the reproduction of echinoids (Kelly, 2001). Bishop (1985) noted that gametogenesis proceeded at temperatures between 11 and 19°C although continued exposure to 19°C destroyed synchronicity of gametogenesis between individuals. Embryos and larvae developed abnormally after 24hr at 15°C (Tyler & Young 1998) but normally at the other temperatures tested (4, 7 and 11°C). Tyler & Young (1998) concluded that embryos and larvae were more tolerant of depth and temperature than adults. Bishop (1985) suggested that Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate and resultant metabolic stress. Therefore, Echinus esculentus is likely to be intolerant of chronic long-term temperature change but would probably be more intolerant of sudden or short-term acute change (e.g. 5°C for one week) in temperature. 

Elevated seawater temperatures generally increase the metabolic rate of bryozoans and are expected to affect calcification (Smith, 2014; Moreno, 2020). The bryozoan Securiflustra securifrons has a northern distribution, recorded at temperatures from -5-15°C (www.obis.org). Therefore, Securiflustra securifrons has is highly likely to be affected by long-term changes in temperature. However, the bryozoan Parasmittina trispinosa has been recorded from Britain, north to western Norway, the Faroe Isles, north western Atlantic coasts, the Californian coast and the Gulf of Mexico. Parasmittina trispinosa generally occurs in temperatures between 5 and 15°C, however, there are records of this species between 25 and 30°C (www.obis.org). Therefore, Parasmittina trispinosa is considered unlikely to be affected by long-term changes in temperature.

Cocito & Sgorbini (2014) studied spatial and temporal patterns of colonial bryozoans in the Ligurian Sea for over nine years. The warming event (a temperature of 23.87 ± 1.4°C at 11 m and of 22.27 ± 1.2°C at 22 m) in the eastern Ligurian Sea (NW Mediterranean) in 1999 caused rapid declines in the abundance of the bryozoan Pentapora between 11 and 22 m depth with an 86% reduction in colony cover (Cocito & Sgorbini, 2014). Recovery was gradual, with communities at 22 m deep recovering to pre-disturbance levels within four years, however, none of the larger colonies (>1,000 cm²) at 11 m deep survived after the first disturbance.

Spirobranchus triqueter occurs as far south as the Mediterranean. Therefore, the species will be subject to a wider range of temperatures than experienced in the British Isles (www.obis.org). Castric-Fey (1983) found that animals settling during spring showed the best growth rate and the best larval settlement occurred in the summer months. Therefore, it is assumed that Spirobranchus triqueter has some tolerance to increased temperatures. 

Sensitivity assessment.  Under the middle, high and extreme emission scenarios seawater temperatures are expected to rise by 3-5°C, with potential southern summer temperatures of 22-24°C. The distribution of Alcyonium digitatum and Securiflustra securifrons suggests that these species may be impacted by ocean warming, as populations of these species are reported where seawater temperatures range up to 20°C and 15°C respectively (www.obis.org).  Caryophyllia smithii, Spirobranchus triqueter and Parasmittina trispinosa are found in geographical locations with higher temperatures than in the UK, therefore these species are unlikely to be affected by an increase in seawater temperature, as these species are likely to adjust to gradual ocean warming. However, the sea urchin Echinus esculentus cannot tolerate high temperatures for prolonged periods, therefore as CR.MCR.EcCr.FaAlCr.Car is a grazed biotope, loss of urchin grazing may result in loss of the biotope and its replacement by another biotope with a more abundant and diverse faunal turf, e.g. CTub.Adig. Therefore, resistance is assessed as ‘Low’ and resilience as ‘Very low’ so that sensitivity is, therefore, assessed as ‘High' at the temperature levels predicted for the end of this century.

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. Further detail.

Evidence

Caryophyllia smithii is a temperate cup coral that has a wide distribution from Norway to the Mediterranean and South Africa in the Atlantic, found in both shallow and deep waters, with records of this species between temperatures of 5 and 30°C. Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in seawater temperature. Therefore, unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns given that gamete release is most likely triggered by seasonal temperature increases (Tranter et al., 1982).

Alcyonium digitatum is a boreal species of octocoral recorded along the Atlantic Coasts of Europe from Portugal to Norway and Iceland, and along the North West Atlantic coasts, at sea surface temperatures between 5 and 20°C but mostly between 10 and 15°C (www.obis.org).  Across this latitudinal gradient species are likely to experience a range of temperatures from approx. 5 and 18°C (Sea temperature, 2015). 

Alcyonium digitatum spawns during the winter months (Hartnoll, 1975). Gametes are fertilized while in the water column and the embryos give rise to actively swimming lecithotrophic planulae that may have an extended pelagic life before they eventually settle (usually within one or two further days) and metamorphose to polyps (Matthews, 1917; Hartnoll, 1975). However, laboratory experiments have observed settlement failure to occur in unsuitable conditions (Hartnoll, 1975). As spawning occurs when sea temperatures are low there is the probability that spawning and settlement could be correlated to climatic conditions, therefore global warming could impact the reproduction and recruitment of Alcyonium digitatum. However, the combination of spawning in winter and the long pelagic lifespan may allow a considerable length of time for the planulae to disperse, settle and metamorphose ahead of the spring plankton bloom (Hartnoll, 1975).

The duration, dispersal and survival of planktonic larvae are dependent on several factors including temperature (O’Connor et al., 2007). O’Connor et al. (2007) reported planktonic larval duration to increase with temperature, therefore cold-water species could see an increase in planktonic larval duration under global warming trends. Larval survival has been reported to decrease exponentially with time (planktonic larval duration) (O’Connor et al., 2007). Elevated temperatures may increase the occurrence of octocoral diseases caused by pathogens that act opportunistically to attack hosts that are under stressful conditions. For example, Cerrano et al. (2000) reported that ecosystems in the Mediterranean are rapidly declining from extensive attacks by microorganisms correlating to elevated seawater temperatures. 

Echinus esculentus is a sea urchin distributed across the N.E. Atlantic from Iceland, north to Finmark, Norway and south to Portugal. Echinus esculentus is common on most coasts of the British Isles but absent from most of the east coast of England, the eastern English Channel and some parts of north Wales. Echinus esculentus has been recorded primarily between sea temperatures of 5 and 15°C (www.obis.org). 

Echinus esculentus occurred at temperatures between 0 and 18°C in Limfjord, Denmark (Ursin 1960). Temperature, photoperiod and food availability are considered to be factors that control the reproduction of echinoids (Kelly, 2001). Bishop (1985) noted that gametogenesis proceeded at temperatures between 11 and 19°C although continued exposure to 19°C destroyed synchronicity of gametogenesis between individuals. Embryos and larvae developed abnormally after 24hr at 15°C (Tyler & Young 1998) but normally at the other temperatures tested (4, 7 and 11°C). Tyler & Young (1998) concluded that embryos and larvae were more tolerant of depth and temperature than adults. Bishop (1985) suggested that Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate and resultant metabolic stress. Therefore, Echinus esculentus is likely to be intolerant of chronic long-term temperature change but would probably be more intolerant of sudden or short-term acute change (e.g. 5°C for one week) in temperature. 

Elevated seawater temperatures generally increase the metabolic rate of bryozoans and are expected to affect calcification (Smith, 2014; Moreno, 2020). The bryozoan Securiflustra securifrons has a northern distribution, recorded at temperatures from -5-15°C (www.obis.org). Therefore, Securiflustra securifrons has is highly likely to be affected by long-term changes in temperature. However, the bryozoan Parasmittina trispinosa has been recorded from Britain, north to western Norway, the Faroe Isles, north western Atlantic coasts, the Californian coast and the Gulf of Mexico. Parasmittina trispinosa generally occurs in temperatures between 5 and 15°C, however, there are records of this species between 25 and 30°C (www.obis.org). Therefore, Parasmittina trispinosa is considered unlikely to be affected by long-term changes in temperature.

Cocito & Sgorbini (2014) studied spatial and temporal patterns of colonial bryozoans in the Ligurian Sea for over nine years. The warming event (a temperature of 23.87 ± 1.4°C at 11 m and of 22.27 ± 1.2°C at 22 m) in the eastern Ligurian Sea (NW Mediterranean) in 1999 caused rapid declines in the abundance of the bryozoan Pentapora between 11 and 22 m depth with an 86% reduction in colony cover (Cocito & Sgorbini, 2014). Recovery was gradual, with communities at 22 m deep recovering to pre-disturbance levels within four years, however, none of the larger colonies (>1,000 cm²) at 11 m deep survived after the first disturbance.

Spirobranchus triqueter occurs as far south as the Mediterranean. Therefore, the species will be subject to a wider range of temperatures than experienced in the British Isles (www.obis.org). Castric-Fey (1983) found that animals settling during spring showed the best growth rate and the best larval settlement occurred in the summer months. Therefore, it is assumed that Spirobranchus triqueter has some tolerance to increased temperatures. 

Sensitivity assessment.  Under the middle, high and extreme emission scenarios seawater temperatures are expected to rise by 3-5°C, with potential southern summer temperatures of 22-24°C. The distribution of Alcyonium digitatum and Securiflustra securifrons suggests that these species may be impacted by ocean warming, as populations of these species are reported where seawater temperatures range up to 20°C and 15°C respectively (www.obis.org).  Caryophyllia smithii, Spirobranchus triqueter and Parasmittina trispinosa are found in geographical locations with higher temperatures than in the UK, therefore these species are unlikely to be affected by an increase in seawater temperature, as these species are likely to adjust to gradual ocean warming. However, the sea urchin Echinus esculentus cannot tolerate high temperatures for prolonged periods, therefore as CR.MCR.EcCr.FaAlCr.Car is a grazed biotope, loss of urchin grazing may result in loss of the biotope and its replacement by another biotope with a more abundant and diverse faunal turf, e.g. CTub.Adig. Therefore, resistance is assessed as ‘Low’ and resilience as ‘Very low’ so that sensitivity is, therefore, assessed as ‘High' at the temperature levels predicted for the end of this century.

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. Further detail.

Evidence

Caryophyllia smithii is a temperate cup coral that has a wide distribution from Norway to the Mediterranean and South Africa in the Atlantic, found in both shallow and deep waters, with records of this species between temperatures of 5 and 30°C. Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in seawater temperature. Therefore, unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns given that gamete release is most likely triggered by seasonal temperature increases (Tranter et al., 1982).

Alcyonium digitatum is a boreal species of octocoral recorded along the Atlantic Coasts of Europe from Portugal to Norway and Iceland, and along the North West Atlantic coasts, at sea surface temperatures between 5 and 20°C but mostly between 10 and 15°C (www.obis.org).  Across this latitudinal gradient species are likely to experience a range of temperatures from approx. 5 and 18°C (Sea temperature, 2015). 

Alcyonium digitatum spawns during the winter months (Hartnoll, 1975). Gametes are fertilized while in the water column and the embryos give rise to actively swimming lecithotrophic planulae that may have an extended pelagic life before they eventually settle (usually within one or two further days) and metamorphose to polyps (Matthews, 1917; Hartnoll, 1975). However, laboratory experiments have observed settlement failure to occur in unsuitable conditions (Hartnoll, 1975). As spawning occurs when sea temperatures are low there is the probability that spawning and settlement could be correlated to climatic conditions, therefore global warming could impact the reproduction and recruitment of Alcyonium digitatum. However, the combination of spawning in winter and the long pelagic lifespan may allow a considerable length of time for the planulae to disperse, settle and metamorphose ahead of the spring plankton bloom (Hartnoll, 1975).

The duration, dispersal and survival of planktonic larvae are dependent on several factors including temperature (O’Connor et al., 2007). O’Connor et al. (2007) reported planktonic larval duration to increase with temperature, therefore cold-water species could see an increase in planktonic larval duration under global warming trends. Larval survival has been reported to decrease exponentially with time (planktonic larval duration) (O’Connor et al., 2007). Elevated temperatures may increase the occurrence of octocoral diseases caused by pathogens that act opportunistically to attack hosts that are under stressful conditions. For example, Cerrano et al. (2000) reported that ecosystems in the Mediterranean are rapidly declining from extensive attacks by microorganisms correlating to elevated seawater temperatures. 

Echinus esculentus is a sea urchin distributed across the N.E. Atlantic from Iceland, north to Finmark, Norway and south to Portugal. Echinus esculentus is common on most coasts of the British Isles but absent from most of the east coast of England, the eastern English Channel and some parts of north Wales. Echinus esculentus has been recorded primarily between sea temperatures of 5 and 15°C (www.obis.org). 

Echinus esculentus occurred at temperatures between 0 and 18°C in Limfjord, Denmark (Ursin 1960). Temperature, photoperiod and food availability are considered to be factors that control the reproduction of echinoids (Kelly, 2001). Bishop (1985) noted that gametogenesis proceeded at temperatures between 11 and 19°C although continued exposure to 19°C destroyed synchronicity of gametogenesis between individuals. Embryos and larvae developed abnormally after 24hr at 15°C (Tyler & Young 1998) but normally at the other temperatures tested (4, 7 and 11°C). Tyler & Young (1998) concluded that embryos and larvae were more tolerant of depth and temperature than adults. Bishop (1985) suggested that Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate and resultant metabolic stress. Therefore, Echinus esculentus is likely to be intolerant of chronic long-term temperature change but would probably be more intolerant of sudden or short-term acute change (e.g. 5°C for one week) in temperature. 

Elevated seawater temperatures generally increase the metabolic rate of bryozoans and are expected to affect calcification (Smith, 2014; Moreno, 2020). The bryozoan Securiflustra securifrons has a northern distribution, recorded at temperatures from -5-15°C (www.obis.org). Therefore, Securiflustra securifrons has is highly likely to be affected by long-term changes in temperature. However, the bryozoan Parasmittina trispinosa has been recorded from Britain, north to western Norway, the Faroe Isles, north western Atlantic coasts, the Californian coast and the Gulf of Mexico. Parasmittina trispinosa generally occurs in temperatures between 5 and 15°C, however, there are records of this species between 25 and 30°C (www.obis.org). Therefore, Parasmittina trispinosa is considered unlikely to be affected by long-term changes in temperature.

Cocito & Sgorbini (2014) studied spatial and temporal patterns of colonial bryozoans in the Ligurian Sea for over nine years. The warming event (a temperature of 23.87 ± 1.4°C at 11 m and of 22.27 ± 1.2°C at 22 m) in the eastern Ligurian Sea (NW Mediterranean) in 1999 caused rapid declines in the abundance of the bryozoan Pentapora between 11 and 22 m depth with an 86% reduction in colony cover (Cocito & Sgorbini, 2014). Recovery was gradual, with communities at 22 m deep recovering to pre-disturbance levels within four years, however, none of the larger colonies (>1,000 cm²) at 11 m deep survived after the first disturbance.

Spirobranchus triqueter occurs as far south as the Mediterranean. Therefore, the species will be subject to a wider range of temperatures than experienced in the British Isles (www.obis.org). Castric-Fey (1983) found that animals settling during spring showed the best growth rate and the best larval settlement occurred in the summer months. Therefore, it is assumed that Spirobranchus triqueter has some tolerance to increased temperatures. 

Sensitivity assessment.  Under the middle, high and extreme emission scenarios seawater temperatures are expected to rise by 3-5°C, with potential southern summer temperatures of 22-24°C. The distribution of Alcyonium digitatum and Securiflustra securifrons suggests that these species may be impacted by ocean warming, as populations of these species are reported where seawater temperatures range up to 20°C and 15°C respectively (www.obis.org).  Caryophyllia smithii, Spirobranchus triqueter and Parasmittina trispinosa are found in geographical locations with higher temperatures than in the UK, therefore these species are unlikely to be affected by an increase in seawater temperature, as these species are likely to adjust to gradual ocean warming. However, the sea urchin Echinus esculentus cannot tolerate high temperatures for prolonged periods, therefore as CR.MCR.EcCr.FaAlCr.Car is a grazed biotope, loss of urchin grazing may result in loss of the biotope and its replacement by another biotope with a more abundant and diverse faunal turf, e.g. CTub.Adig. Therefore, resistance is assessed as ‘Low’ and resilience as ‘Very low’ so that sensitivity is, therefore, assessed as ‘High' at the temperature levels predicted for the end of this century.

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. Further detail.

Evidence

Marine heatwaves are extreme weather events defined as periods of extreme sea surface temperature that persists for days to months (Frölicher et al., 2018). Marine heatwaves are predicted to occur more frequently, last for longer and at increased intensity by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018).

Caryophyllia smithii is a temperate cup coral that has a wide distribution and is found in both shallow and deep waters, with records of this species between temperatures of 5 and 30°C. Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in seawater temperature. Therefore, unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns given that gamete release is most likely triggered by seasonal temperature increases (Tranter et al., 1982). Therefore, marine heatwaves could impact the reproduction and recruitment of Caryophyllia smithii. 

No evidence on the impacts of marine heatwaves on Alcyonium digitatum was found, this species appears to be restricted to colder waters. With Alcyonium digitatum occurring in seawater temperatures between 5 and 20°C (www.obis.org). Therefore, Alcyonium digitatum is likely to be impacted by heatwaves under both emission scenarios. Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate resulting in metabolic stress (Bishop 1985). Bishop (1985) observed gametogenesis to occur between 11 and 19°C however, continued exposure to 19°C disrupted gametogenesis. In addition, embryos and larvae developed abnormally after 24hr exposure to 15°C (Bishop, 1985). Therefore, marine heatwaves have the potential to impact the reproduction, recruitment and survival of Echinus esculentus. 

Elevated seawater temperatures generally increase the metabolic rate of bryozoans and are expected to affect calcification (Smith, 2014; Leveroni, 2020). No evidence on the impacts of marine heatwaves on Securiflustra securifrons was found, however as this species occurs between -5-15°C (www.obis.org), therefore there is a high chance of the species being affected by heatwaves. Similarly, no evidence on the impacts of marine heatwaves on Parasmittina trispinosa was found. However, although Parasmittina trispinosa generally occurs in temperatures between 5 and 15°C, there are records of this species between 25 and 30°C (www.obis.org). Therefore, Parasmittina trispinosa is considered unlikely to be affected by long-term changes in temperature in the UK, as Parasmittina trispinosa is likely to acclimate to temperatures with time. However, the occurrence of marine heatwaves could cause mass mortality to populations that have not been acclimated to warmer temperatures.

Cocito & Sgorbini (2014) studied spatial and temporal patterns of colonial bryozoans in the Ligurian Sea for over nine years. The warming event (a temperature of 23.87 ± 1.4°C at 11 m and of 22.27 ± 1.2°C at 22 m) in the eastern Ligurian Sea (NW Mediterranean) in 1999 caused rapid declines in the abundance of the bryozoan Pentapora between 11 and 22 m depth with an 86% reduction in colony cover (Cocito & Sgorbini, 2014). Recovery was gradual, with communities at 22 m deep recovering to pre-disturbance levels within four years, however, none of the larger colonies (>1,000 cm²) at 11 m deep survived after the first disturbance. Spirobranchus triqueter is recorded as abundant from the Iberian Peninsula, Spain (Ramos, 2010) as well as from the Alexandria coast of Egypt, Mediterranean Sea (Sarah, 2010). Across this latitudinal gradient, Spirobranchus triqueter is likely to experience a range of temperatures from approximately 5-28°C (Sea temperature, 2015), and is therefore unlikely to be affected by marine heatwaves. 

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 occurred 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. There is no experimental evidence of the impact of marine heatwaves on the characteristic species of this biotope, however as CR.MCR.EcCr.FaAlCr.Car is a grazed biotope and the primary grazer Echinus esculentus cannot tolerate high temperatures for prolonged periods, under the middle and high emissions scenario resistance, is assessed as “Low”, and resilience is assessed as “Low”, so the biotope is assessed as “High” sensitivity.

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. Further detail.

Evidence

Marine heatwaves are extreme weather events defined as periods of extreme sea surface temperature that persists for days to months (Frölicher et al., 2018). Marine heatwaves are predicted to occur more frequently, last for longer and at increased intensity by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018).

Caryophyllia smithii is a temperate cup coral that has a wide distribution and is found in both shallow and deep waters, with records of this species between temperatures of 5 and 30°C. Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in seawater temperature. Therefore, unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns given that gamete release is most likely triggered by seasonal temperature increases (Tranter et al., 1982). Therefore, marine heatwaves could impact the reproduction and recruitment of Caryophyllia smithii. 

No evidence on the impacts of marine heatwaves on Alcyonium digitatum was found, this species appears to be restricted to colder waters. With Alcyonium digitatum occurring in seawater temperatures between 5 and 20°C (www.obis.org). Therefore, Alcyonium digitatum is likely to be impacted by heatwaves under both emission scenarios. Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate resulting in metabolic stress (Bishop 1985). Bishop (1985) observed gametogenesis to occur between 11 and 19°C however, continued exposure to 19°C disrupted gametogenesis. In addition, embryos and larvae developed abnormally after 24hr exposure to 15°C (Bishop, 1985). Therefore, marine heatwaves have the potential to impact the reproduction, recruitment and survival of Echinus esculentus. 

Elevated seawater temperatures generally increase the metabolic rate of bryozoans and are expected to affect calcification (Smith, 2014; Leveroni, 2020). No evidence on the impacts of marine heatwaves on Securiflustra securifrons was found, however as this species occurs between -5-15°C (www.obis.org), therefore there is a high chance of the species being affected by heatwaves. Similarly, no evidence on the impacts of marine heatwaves on Parasmittina trispinosa was found. However, although Parasmittina trispinosa generally occurs in temperatures between 5 and 15°C, there are records of this species between 25 and 30°C (www.obis.org). Therefore, Parasmittina trispinosa is considered unlikely to be affected by long-term changes in temperature in the UK, as Parasmittina trispinosa is likely to acclimate to temperatures with time. However, the occurrence of marine heatwaves could cause mass mortality to populations that have not been acclimated to warmer temperatures.

Cocito & Sgorbini (2014) studied spatial and temporal patterns of colonial bryozoans in the Ligurian Sea for over nine years. The warming event (a temperature of 23.87 ± 1.4°C at 11 m and of 22.27 ± 1.2°C at 22 m) in the eastern Ligurian Sea (NW Mediterranean) in 1999 caused rapid declines in the abundance of the bryozoan Pentapora between 11 and 22 m depth with an 86% reduction in colony cover (Cocito & Sgorbini, 2014). Recovery was gradual, with communities at 22 m deep recovering to pre-disturbance levels within four years, however, none of the larger colonies (>1,000 cm²) at 11 m deep survived after the first disturbance. Spirobranchus triqueter is recorded as abundant from the Iberian Peninsula, Spain (Ramos, 2010) as well as from the Alexandria coast of Egypt, Mediterranean Sea (Sarah, 2010). Across this latitudinal gradient, Spirobranchus triqueter is likely to experience a range of temperatures from approximately 5-28°C (Sea temperature, 2015), and is therefore unlikely to be affected by marine heatwaves. 

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 occurred 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. There is no experimental evidence of the impact of marine heatwaves on the characteristic species of this biotope, however as CR.MCR.EcCr.FaAlCr.Car is a grazed biotope and the primary grazer Echinus esculentus cannot tolerate high temperatures for prolonged periods, under the middle and high emissions scenario resistance, is assessed as “Low”, and resilience is assessed as “Low”, so the biotope is assessed as “High” sensitivity.

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. Further detail 

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). In general, it is thought that calcifying invertebrates will be more sensitive to ocean acidification than non-calcifying invertebrates, which appear to have a more mixed response (Hofmann et al., 2010). It must be noted that many species show variation in their response to pCO₂ independent of their taxonomic group or habitat preferences (Widdicombe & Spicer, 2008; Kroeker et al., 2013).

Ocean acidification has negative impacts on numerous species of coral; however, laboratory evidence has shown that the temperate cup coral Caryophyllia smithii might have some resistance to ocean acidification. Rodolfo-Metalpa et al. (2015) exposed Caryophyllia smithii samples to elevated CO₂ conditions expected for the end of this century for several months. All of the corals survived the treatment and no significant differences in respiration or gross and net calcification rates were observed under high seawater pCO₂. 

No evidence of the impacts of ocean acidification on Alcyonium digitatum was found. However, studies on the impacts of ocean acidification on octocorals reported various responses (Conci et al., 2021). Gomez et al. (2014) found a significant negative correlation between calcification and CO₂ concentrations for Eunicea flexuosa at a pH range of 8.1–7.1. But another study on Eunicea flexuosa observed no significant differences in branch extension and sclerite structure at pH 7.75 and suggested that Eunicea flexuosa had a degree of resilience to ocean acidification (Enochs et al., 2016). Similarly, ocean acidification did not significantly impact the octocorals Ovabunda macrospiculata, Heteroxenia fuscescens and Sarcophyton sp. with no effects on polyp weight and protein concentration, nor any significant differences in chlorophyll abundance or density of zooxanthellae at pH 7.6 and 7.3 when compared to controls at pH 8.2. The findings suggested that the octocoral’s tissue may provide a protective role against acidification (Gabay et al., 2013; Gabay et al., 2014).

The planktonic larval stage is often thought to be the most sensitive stage to ocean acidification in benthic organisms (Kurihara, 2008, Chan et al., 2015).  The embryos of Alcyonium digitatum are neutrally buoyant and float freely for several days before they give rise to actively swimming lecithotrophic planulae which may have an extended pelagic life before they eventually settle (usually within one or two additional days) and metamorphose to polyps (Matthews, 1917; Hartnoll, 1975; Budd, 2008). In laboratory experiments, larvae of Alcyonium digitatum failed to settle within ten days, presumably finding the conditions unsuitable (Hartnoll, 1975), however, the water conditions were not recorded. 

Dupont et al. (2010) analysed the literature and suggested that echinoderms were generally robust to ocean acidification, although different life stages and species were affected differently. Limited evidence on the impacts of ocean acidification on Echinus esculentus was found. However, near future CO₂-driven ocean acidification (-0.4 units for the end of this century) had negative impacts on survival and developmental dynamics of Echinus esculentus (Dupont and Thorndyke, personal communication, 2009).  Evidence on the reproduction or early life stages of Echinus esculentus was not found, however, studies have found a variety of responses to ocean acidification depending on the species of sea urchin. Dworjanyn & Byrne (2018) found acidification to decrease the gonad index of Tripneustes gratilla, with almost no gonads in urchins at pH 7.6 regardless of temperature. Clark et al, (2009) observed the effects of lowered pH on larvae from tropical (Tripneustes gratilla), temperate (Pseudechinus huttoni, Evechinus chloroticus), and a polar species (Sterechinus neumayeri) of sea urchin. The results indicated that the survival of larvae may not be directly affected by the pH levels predicted for 2100, but the low pH may cause reduced growth and calcification, which could compromise survival. Lee et al. (2019) observed metabolic rates of Strongylocentrotus purpuratus larvae to increase with decreasing pH and reach a threshold between pH 7.0 and pH 7.3 where metabolic rates decreased again. Therefore, ocean acidification could have detrimental effects on the survival, reproduction and recruitment of Echinus esculentus. 

Nevertheless, Suckling et al. (2014) emphasized that studies that presented stressors in a shock-type exposure (as above) may reflect stress response outcomes rather than the results of gradual change in the climate. Cross generation echinoderm studies observed a variety of responses to the progeny produced by adults that have been exposed to low pH. The evidence indicated that the effect on progeny depended on the level of acidification and the conditioning duration of the parents (Byrne et al., 2019). Suckling et al. (2014) found that when Psammechinus miliaris larvae were raised from parents pre-exposed to low pH conditions (pH 7.7 compared to control pH of 7.98), settlement rates were similar to control larvae, and the test (i.e. the urchin shell) diameter was larger, which suggested that this species can acclimate and possibly adapt to low pH conditions. Similarly, Clark et al. (2019) observed that gene expression profiles associated with transgenerational plasticity contributed to Psammechinus miliaris larval resilience when the adults were conditioned to low pH.

From observations at natural vent sites, Connell et al. (2018) observed that increased CO₂ enrichment reduced the abundance and feeding rates of primary grazers (urchins, Evechinus chloroticus), allowing turf algae to increase in abundance. Therefore, ocean acidification could cause changes to community structure. 

Bryozoans are invertebrate calcifiers, therefore, they are potentially highly sensitive to ocean acidification (Smith, 2009). The decrease in water pH from global climate change could cause corrosion, changes in mineralogy and decrease the survival of bryozoans (Smith, 2014). No evidence on the impacts of ocean acidification on the characterizing bryozoan species Parasmittina trispinosa was found. However, Swezey et al. (2017) observed that populations of bryozoans raised under high CO₂ (1254 μatm; pH 7.60) conditions grew faster, invested less in reproduction and produced lighter skeletons when compared to genetically identical clones raised under current surface atmospheric CO₂ values (400 μatm; pH 8.04). In addition, the bryozoans under high CO₂ altered the Mg/Ca ratio of skeletal calcite, which could be a protective mechanism against acidification (Swezey et al., 2017). 

Lombardi et al. (2011) investigated the impacts of ocean acidification on the growth, organic tissue and protein profile of bryozoan Myriapora truncata along a gradient of different pH levels in a natural volcanic CO2 vent site. At sites with normal pH levels (mean pH 8.10), Myriapora truncata produced new and complete zooids. However, at the intermediate (pH 7.83) and low pH (pH 7.32) sites neither partial nor complete zooids were produced. At the intermediate pH sites, Myriapora truncata increased its skeleton thickness suggesting a protective defence against dissolution, but at the low pH sites, there was a decrease in skeletal weights and corrosion of skeletal structures. Additionally, at intermediate and low pH sites Myriapora truncata upregulated protein production to potentially overcome the low pH conditions, however, the upregulation came at a cost, and fitness was reduced resulting in mortality particularly in the lower pH sites. 

Studies on the impacts of ocean acidification on the calcareous tube structure of serpulidae polychaete worms observed reductions of tube elongation, elasticity and strength when exposed to reduced pH conditions (Chan et al.,2012; Li et al., 2014; Dıáz-Castanẽda et al., 2019). Dıáz-Castanẽda et al (2019). observed pH to effect trochophore size and post-settlement tube growth, with post-settlement tubes half the size of those at current ocean pH levels. Li et al (2014) observed changes to the structure, volume and density of the tubes, with reductions in hardiness and elasticity, in addition to a 64% reduction in tube crushing force. The reduction in tube size and hardiness could impact the survival of the species with increased predation and reduction in the ability to withstand wave force (Chan et al., 2012; Li et al., 2014). 

Sensitivity Assessment. Laboratory evidence has shown that Caryophyllia smithii has some resistance to ocean acidification, therefore the evidence suggests that Caryophyllia smithii may be able to tolerate future pH predictions, however, impacts on reproduction and recruitment were not reviewed.  While limited evidence of the impacts of ocean acidification on Echinus esculentus was found, the effects of ocean acidification on other echinoderms species have shown species dependent sensitivity. No evidence of the effect of ocean acidification on Alcyonium digitatum was found, although the effects of ocean acidification on other species of octocoral have indicated some resilience to low pH. Though Alcyonium digitatum larvae settlement has been reported to be sensitive to environmental conditions, however, the conditions were not verified. Bryozoans appear to be highly sensitive to ocean acidification, with impacts on health, survival and reproduction.  Unfortunately, at present, there are no studies to determine whether Echinus esculentus can adapt or acclimate to future pH conditions, but on the evidence available, Echinus esculentus could be lost from this biotope under the predicted emissions scenarios. As CR.MCR.EcCr.FaAlCr.Car is a grazed biotope, loss of urchin grazing may result in loss of the biotope and its replacement by another biotope with a more abundant and diverse faunal turf, e.g. CTub.Adig. Therefore, under both the middle and high emission scenarios (0.15 and 0.35 pH unit decrease, respectively) the biotope is assessed as having a resistance level of ‘Low’, and a resilience level of ‘Very low’ because of the long-term nature of ocean acidification, leading to an assessment of ‘High’ sensitivity at the benchmark level, albeit with ‘Low’ confidence.

Ocean acidification (middle)

Middle emission scenario benchmark: a further decrease in pH of 0.15 (annual mean) and 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. Further detail.

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). In general, it is thought that calcifying invertebrates will be more sensitive to ocean acidification than non-calcifying invertebrates, which appear to have a more mixed response (Hofmann et al., 2010). It must be noted that many species show variation in their response to pCO₂ independent of their taxonomic group or habitat preferences (Widdicombe & Spicer, 2008; Kroeker et al., 2013).

Ocean acidification has negative impacts on numerous species of coral; however, laboratory evidence has shown that the temperate cup coral Caryophyllia smithii might have some resistance to ocean acidification. Rodolfo-Metalpa et al. (2015) exposed Caryophyllia smithii samples to elevated CO₂ conditions expected for the end of this century for several months. All of the corals survived the treatment and no significant differences in respiration or gross and net calcification rates were observed under high seawater pCO₂. 

No evidence of the impacts of ocean acidification on Alcyonium digitatum was found. However, studies on the impacts of ocean acidification on octocorals reported various responses (Conci et al., 2021). Gomez et al. (2014) found a significant negative correlation between calcification and CO₂ concentrations for Eunicea flexuosa at a pH range of 8.1–7.1. But another study on Eunicea flexuosa observed no significant differences in branch extension and sclerite structure at pH 7.75 and suggested that Eunicea flexuosa had a degree of resilience to ocean acidification (Enochs et al., 2016). Similarly, ocean acidification did not significantly impact the octocorals Ovabunda macrospiculata, Heteroxenia fuscescens and Sarcophyton sp. with no effects on polyp weight and protein concentration, nor any significant differences in chlorophyll abundance or density of zooxanthellae at pH 7.6 and 7.3 when compared to controls at pH 8.2. The findings suggested that the octocoral’s tissue may provide a protective role against acidification (Gabay et al., 2013; Gabay et al., 2014).

The planktonic larval stage is often thought to be the most sensitive stage to ocean acidification in benthic organisms (Kurihara, 2008, Chan et al., 2015).  The embryos of Alcyonium digitatum are neutrally buoyant and float freely for several days before they give rise to actively swimming lecithotrophic planulae which may have an extended pelagic life before they eventually settle (usually within one or two additional days) and metamorphose to polyps (Matthews, 1917; Hartnoll, 1975; Budd, 2008). In laboratory experiments, larvae of Alcyonium digitatum failed to settle within ten days, presumably finding the conditions unsuitable (Hartnoll, 1975), however, the water conditions were not recorded. 

Dupont et al. (2010) analysed the literature and suggested that echinoderms were generally robust to ocean acidification, although different life stages and species were affected differently. Limited evidence on the impacts of ocean acidification on Echinus esculentus was found. However, near future CO₂-driven ocean acidification (-0.4 units for the end of this century) had negative impacts on survival and developmental dynamics of Echinus esculentus (Dupont and Thorndyke, personal communication, 2009).  Evidence on the reproduction or early life stages of Echinus esculentus was not found, however, studies have found a variety of responses to ocean acidification depending on the species of sea urchin. Dworjanyn & Byrne (2018) found acidification to decrease the gonad index of Tripneustes gratilla, with almost no gonads in urchins at pH 7.6 regardless of temperature. Clark et al, (2009) observed the effects of lowered pH on larvae from tropical (Tripneustes gratilla), temperate (Pseudechinus huttoni, Evechinus chloroticus), and a polar species (Sterechinus neumayeri) of sea urchin. The results indicated that the survival of larvae may not be directly affected by the pH levels predicted for 2100, but the low pH may cause reduced growth and calcification, which could compromise survival. Lee et al. (2019) observed metabolic rates of Strongylocentrotus purpuratus larvae to increase with decreasing pH and reach a threshold between pH 7.0 and pH 7.3 where metabolic rates decreased again. Therefore, ocean acidification could have detrimental effects on the survival, reproduction and recruitment of Echinus esculentus. 

Nevertheless, Suckling et al. (2014) emphasized that studies that presented stressors in a shock-type exposure (as above) may reflect stress response outcomes rather than the results of gradual change in the climate. Cross generation echinoderm studies observed a variety of responses to the progeny produced by adults that have been exposed to low pH. The evidence indicated that the effect on progeny depended on the level of acidification and the conditioning duration of the parents (Byrne et al., 2019). Suckling et al. (2014) found that when Psammechinus miliaris larvae were raised from parents pre-exposed to low pH conditions (pH 7.7 compared to control pH of 7.98), settlement rates were similar to control larvae, and the test (i.e. the urchin shell) diameter was larger, which suggested that this species can acclimate and possibly adapt to low pH conditions. Similarly, Clark et al. (2019) observed that gene expression profiles associated with transgenerational plasticity contributed to Psammechinus miliaris larval resilience when the adults were conditioned to low pH.

From observations at natural vent sites, Connell et al. (2018) observed that increased CO₂ enrichment reduced the abundance and feeding rates of primary grazers (urchins, Evechinus chloroticus), allowing turf algae to increase in abundance. Therefore, ocean acidification could cause changes to community structure. 

Bryozoans are invertebrate calcifiers, therefore, they are potentially highly sensitive to ocean acidification (Smith, 2009). The decrease in water pH from global climate change could cause corrosion, changes in mineralogy and decrease the survival of bryozoans (Smith, 2014). No evidence on the impacts of ocean acidification on the characterizing bryozoan species Parasmittina trispinosa was found. However, Swezey et al. (2017) observed that populations of bryozoans raised under high CO₂ (1254 μatm; pH 7.60) conditions grew faster, invested less in reproduction and produced lighter skeletons when compared to genetically identical clones raised under current surface atmospheric CO₂ values (400 μatm; pH 8.04). In addition, the bryozoans under high CO₂ altered the Mg/Ca ratio of skeletal calcite, which could be a protective mechanism against acidification (Swezey et al., 2017). 

Lombardi et al. (2011) investigated the impacts of ocean acidification on the growth, organic tissue and protein profile of bryozoan Myriapora truncata along a gradient of different pH levels in a natural volcanic CO2 vent site. At sites with normal pH levels (mean pH 8.10), Myriapora truncata produced new and complete zooids. However, at the intermediate (pH 7.83) and low pH (pH 7.32) sites neither partial nor complete zooids were produced. At the intermediate pH sites, Myriapora truncata increased its skeleton thickness suggesting a protective defence against dissolution, but at the low pH sites, there was a decrease in skeletal weights and corrosion of skeletal structures. Additionally, at intermediate and low pH sites Myriapora truncata upregulated protein production to potentially overcome the low pH conditions, however, the upregulation came at a cost, and fitness was reduced resulting in mortality particularly in the lower pH sites. 

Studies on the impacts of ocean acidification on the calcareous tube structure of serpulidae polychaete worms observed reductions of tube elongation, elasticity and strength when exposed to reduced pH conditions (Chan et al.,2012; Li et al., 2014; Dıáz-Castanẽda et al., 2019). Dıáz-Castanẽda et al (2019). observed pH to effect trochophore size and post-settlement tube growth, with post-settlement tubes half the size of those at current ocean pH levels. Li et al (2014) observed changes to the structure, volume and density of the tubes, with reductions in hardiness and elasticity, in addition to a 64% reduction in tube crushing force. The reduction in tube size and hardiness could impact the survival of the species with increased predation and reduction in the ability to withstand wave force (Chan et al., 2012; Li et al., 2014). 

Sensitivity Assessment. Laboratory evidence has shown that Caryophyllia smithii has some resistance to ocean acidification, therefore the evidence suggests that Caryophyllia smithii may be able to tolerate future pH predictions, however, impacts on reproduction and recruitment were not reviewed.  While limited evidence of the impacts of ocean acidification on Echinus esculentus was found, the effects of ocean acidification on other echinoderms species have shown species dependent sensitivity. No evidence of the effect of ocean acidification on Alcyonium digitatum was found, although the effects of ocean acidification on other species of octocoral have indicated some resilience to low pH. Though Alcyonium digitatum larvae settlement has been reported to be sensitive to environmental conditions, however, the conditions were not verified. Bryozoans appear to be highly sensitive to ocean acidification, with impacts on health, survival and reproduction.  Unfortunately, at present, there are no studies to determine whether Echinus esculentus can adapt or acclimate to future pH conditions, but on the evidence available, Echinus esculentus could be lost from this biotope under the predicted emissions scenarios. As CR.MCR.EcCr.FaAlCr.Car is a grazed biotope, loss of urchin grazing may result in loss of the biotope and its replacement by another biotope with a more abundant and diverse faunal turf, e.g. CTub.Adig. Therefore, under both the middle and high emission scenarios (0.15 and 0.35 pH unit decrease, respectively) the biotope is assessed as having a resistance level of ‘Low’, and a resilience level of ‘Very low’ because of the long-term nature of ocean acidification, leading to an assessment of ‘High’ sensitivity at the benchmark level, albeit with ‘Low’ confidence.

Sea level rise (extreme)

Extreme scenario benchmark: a 107 cm rise in average UK by the end of this century (2018-2100). Further detail.

Evidence

Sea-level rise is occurring through a combination of thermal expansion and ice melt.  Sea levels have risen 1-3 mm/yr. in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). Evidence appears to suggest that impacts of sea-level rise on exposure or tidal energy will be non-linear and site-specific (Pickering et al., 2012, Li et al., 2016).  This biotope occurs on mixed sediment, in moderately exposed to sheltered areas, subject to strong to weak tidal streams and, therefore, should be reasonably robust to any changes which occur. Furthermore, this biotope occurs at depths of 10-30 m around the UK, and sea-level rises predicted for the end of this century should have limited impacts on this biotope.

Sensitivity assessment. As this biotope CR.MCR.EcCr.FaAlCr.Car can occur from 10-30m depth, in a range of different energy environments, it is assumed that a sea-level rise of 50 cm, 70 cm or 107 cm (middle and high emission, and extreme scenarios) would have limited effect. Therefore, resistance is assessed as ‘High’ under all three scenarios, so that resilience is ‘High’ and sensitivity assessed as ‘Not sensitive’. 

Sea level rise (high)

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

Evidence

Sea-level rise is occurring through a combination of thermal expansion and ice melt.  Sea levels have risen 1-3 mm/yr. in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). Evidence appears to suggest that impacts of sea-level rise on exposure or tidal energy will be non-linear and site-specific (Pickering et al., 2012, Li et al., 2016).  This biotope occurs on mixed sediment, in moderately exposed to sheltered areas, subject to strong to weak tidal streams and, therefore, should be reasonably robust to any changes which occur. Furthermore, this biotope occurs at depths of 10-30 m around the UK, and sea-level rises predicted for the end of this century should have limited impacts on this biotope.

Sensitivity assessment. As this biotope CR.MCR.EcCr.FaAlCr.Car can occur from 10-30m depth, in a range of different energy environments, it is assumed that a sea-level rise of 50 cm, 70 cm or 107 cm (middle and high emission, and extreme scenarios) would have limited effect. Therefore, resistance is assessed as ‘High’ under all three scenarios, so that resilience is ‘High’ and sensitivity assessed as ‘Not sensitive’. 

Sea level rise (middle)

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

Evidence

Sea-level rise is occurring through a combination of thermal expansion and ice melt.  Sea levels have risen 1-3 mm/yr. in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). Evidence appears to suggest that impacts of sea-level rise on exposure or tidal energy will be non-linear and site-specific (Pickering et al., 2012, Li et al., 2016).  This biotope occurs on mixed sediment, in moderately exposed to sheltered areas, subject to strong to weak tidal streams and, therefore, should be reasonably robust to any changes which occur. Furthermore, this biotope occurs at depths of 10-30 m around the UK, and sea-level rises predicted for the end of this century should have limited impacts on this biotope.

Sensitivity assessment. As this biotope CR.MCR.EcCr.FaAlCr.Car can occur from 10-30m depth, in a range of different energy environments, it is assumed that a sea-level rise of 50 cm, 70 cm or 107 cm (middle and high emission, and extreme scenarios) would have limited effect. Therefore, resistance is assessed as ‘High’ under all three scenarios, so that resilience is ‘High’ and sensitivity assessed as ‘Not sensitive’. 

Temperature increase (local)

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

Evidence

The majority of the records of CR.MCR.EcCr.FaAlCr.Car occur in west Scotland, but numerous records occur in south-west Ireland and the south-west of England. Sea surface temperature across this distribution ranges from northern to southern Sea Surface Temperature (SST) of 8-16°C in summer and 6-13°C in winter (Beszczynska-Möller & Dye, 2013). Alcyonium digitatum is described as a northern species by Hiscock et al. (2004) but is distributed from Northern Norway (70°N) to Portugal (41°N) (Hartnoll, 1975; Budd, 2008). Securiflustra securifrons is recorded from Kongsfjorden, Svalbard (Gontar et al., 2001) to the Iberian peninsula in both Spain and Portugal (Ramos, 2010). Across this latitudinal gradient both species are likely to experience a range of temperatures from approximately 5-18°C (Seatemperature, 2015). Spirobranchus triqueter is described as a temperate species by Kupriyanova & Badyaev (1998). Spirobranchus triqueter is recorded as abundant in sub-tidal habitats of Trondheimsfjord (63°N) (Kukliński & Barnes, 2008); no survey reports could be found further north. The most southerly records are from the Iberian peninsula, Spain (Ramos, 2010) as well from the Alexandria coast of Egypt, Mediterranean Sea (Dorgham et al., 2013). Across this latitudinal gradient, Spirobranchus triqueter is likely to experience a range of temperatures from approximately 5-28°C (Seatemperature, 2015). Bishop (1985) suggested that Echinus esculentus cannot tolerate high temperatures for prolonged periods due to increased respiration rate and resultant metabolic stress. Ursin (1960) reported Echinus esculentus occurred at temperatures between 0-18°C in Limfjord, Denmark. Bishop (1985) noted that gametogenesis occurred at 11-19°C however, continued exposure to 19°C disrupted gametogenesis. Embryos and larvae developed abnormally after 24hr exposure to 15°C but normally at 4, 7 and 11°C (Tyler & Young, 1998). Mature examples of Caryophyllia smithii are recorded in Greece (Koukouras, 2010), and are, therefore, unlikely to be physically affected at the benchmark. However, Tranter et al. (1982) suggested Caryophyllia smithii reproduction was cued by seasonal increases in temperature. Therefore, unseasonal increases in temperature may disrupt natural reproductive processes and negatively influence recruitment patterns. Mature examples of Caryophyllia smithii were recorded in Greece.

Sensitivity assessment. Spirobranchus triqueter records from the Alexandria coast of Egypt, Mediterranean Sea (Dorgham et al., 2013) indicate the species is unlikely to be affected at the benchmark level. An increase in sea surface temperature of 2°C for a period of 1 year combined with high temperatures may approach the upper temperature threshold of Alcyonium digitatum, Echinus esculentus, and may, therefore, cause minor declines in abundance. There was insufficient evidence to assess the effect of a short-term increase in temperature of 5°C on Alcyonium digitatum however it may disrupt Echinus esculentus spawning in southern examples of this biotope. If reproduction in Caryophyllia smithii was disrupted by a short-term or long-term change in temperature then recruitment would be impaired for the affected period (one year) but recover subsequently as long as adults survived. Therefore, resistance has been assessed as ‘Medium’, resilience as ‘High’ and sensitivity has been assessed as ‘Low’.

Temperature decrease (local)

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

Evidence

The majority of the records of CR.MCR.EcCr.FaAlCr.Car occur in north-west Scotland, but numerous records occur in south-west Ireland and the south-west of England. Sea surface temperature across this distribution ranges from northern to southern Sea Surface Temperature (SST) of 8-16°C in summer and 6-13°C in winter (Beszczynska-Möller & Dye, 2013).

Alcyonium digitatum is described as a northern species by Hiscock et al. (2004) but is distributed from northern Norway (70°N) to Portugal (41°N) (Hartnoll, 1975; Budd, 2008). Across this latitudinal gradient, both species are likely to experience a range of temperatures from approximately 5-18°C. Alcyonium digitatum was also reported to be apparently unaffected by the severe winter of 1962-1963 where air temperature reached -5.8°C (Crisp, 1964). Securiflustra securifrons is recorded from Kongsfjorden, Svalbard (Gontar et al., 2001) to the Iberian peninsula in both Spain and Portugal (Ramos, 2010). Echinus esculentus has been recorded from the Murmansk Coast, Russia. Due to the high latitude at which Echinus esculentus can occur it is unlikely to be affected at the pressure benchmark. 

Spirobranchus triqueter is described as a temperate species by Kupriyanova & Badyaev (1998). Spirobranchus triqueter is recorded as abundant in subtidal habitats of Trondheimsfjord (63°N) (Kukliński & Barnes, 2008); no survey reports could be found further north. Averaged across several years the lowest winter temperature within Trondheimsfjord is 4.9°C (Seatemperature, 2015). Below 7°C Spirobranchus triqueter is unable to build calcareous tubes (Thomas, 1940). Mature adults may survive a decrease at the pressure benchmark, however, larvae may not be able to attach to the substratum (Riley & Ballerstedt, 2005) if a temperature decrease co-occurred with cold winter temperatures in the UK. However, settlement is reportedly low within winter (see resilience section), and therefore the effects on recruitment are likely to be minor.

Caryophyllia smithii has a northern range limit in the Shetland Isles and southern Norway (NBN, 2015), and may, therefore, be negatively affected by cold temperatures in northern examples of this biotope.

Sensitivity assessment. Alcyonium digitatum, Echinus esculentus have northern/boreal distributions and are unlikely to be affected at the benchmark level. Spirobranchus triqueter is unable to build calcareous tubes at low temperatures, however, during winter, this is unlikely to have any significant effects on recruitment. In addition, the depth of the biotope probably protects it from short-term acute decreases in temperature. The important characterizing Caryophyllia smithii is close to its northern distribution limit within the British Isles and a decrease at the benchmark level may result in some mortality in northern examples of the biotope.  Therefore, resistance is, therefore ‘Medium’, resilience is ‘High’ and sensitivity is ‘Low’.

Salinity increase (local)

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

Evidence

Lyster (1965) tested the tolerance of Spirobranchus triqueter larvae to various hyper and hypo salinity treatments. Larvae were placed in cultures ranging from 0-90‰ and notes were made on the time taken for larvae to die or begin displaying abnormal behaviour. Spirobranchus triqueter larvae were tolerant of salinities ranging from 20-50‰, above 50‰ caused high mortality. Spirobranchus triqueter is, therefore, unlikely to be affected at the pressure benchmark.

Echinoderms are generally stenohaline and possess no osmoregulatory organ (Boolootian, 1966). Therefore, an increase in salinity may cause Echinus esculentus mortality. Alcyonium digitatum’ distribution and the depth at which it occurs also suggest it would not likely experience regular salinity fluctuations and therefore tolerate significant increases in salinity. 

Sensitivity assessment. CR.MCR.EcCr.FaAlCr.Car is restricted to full salinity (Connor et al., 2004), therefore, it is likely that an increase in salinity to >40‰ (e.g. from hypersaline, brine effluents) may cause a decline in the abundance of Alcyonium digitatum and Echinus esculentus. A reduction in Echinus grazing is likely to alter the character of the biotope. Therefore, resistance has been assessed as ‘Low’, resilience as ‘High’. Sensitivity has been assessed as ‘Low’. Due to the lack of information regarding salinity effects on Alcyonium digitatum, Echinus esculentus Caryophyllia smithii confidence in this assessment has been assessed as 'Low'. 

Salinity decrease (local)

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

Evidence

Alcyonium digitatum does inhabit situations such as the entrances to sea lochs (Budd, 2008) or the entrances to estuaries (Braber & Borghouts, 1977) where salinity may vary occasionally. Furthermore,  Alcyonium digitatum is found within a number of variable salinity biotopes, e.g. MCR.ByH.Flu.Hocu. However, its distribution and the depth at which it occurs suggest that Alcyonium digitatum would not likely often experience salinity fluctuations and therefore unlikely to survive significant reductions in salinity (Budd, 2008).

Echinoderms are generally unable to tolerate low salinity (stenohaline) and possess no osmoregulatory organ (Boolootian, 1966). At low salinity, urchins gain weight, and the epidermis loses its pigment as patches are destroyed; prolonged exposure is fatal. However, within Echinus esculentus, there is some evidence to suggest intracellular regulation of osmotic pressure due to increased amino acid concentrations. Furthermore, Echinus esculentus is found within a number of variable and reduced salinity biotopes, e.g. IR.LIR.KVS.SlatPsaVS.

Lyster (1965) tested the tolerance of Spirobranchus triqueter larvae to various hyper and hypo salinity treatments. Larvae were placed in cultures ranging from 0-90‰ and notes were made on the time taken for larvae to die or begin displaying abnormal behaviour. Spirobranchus triqueter larvae can survive very well in salinities down to 20‰, and can tolerate salinities down to 10‰. Adults are tolerant of salinities as low as 3‰, and can be found in areas were salinity ranges from 18-23‰ (Alexander et al., 1935). No information on the salinity tolerance of Caryophyllia smithii was found. However, it is recorded from the CR.LCR.BrAs.NovPro.VS biotope that occurs in 'variable', 'reduced' and 'low' salinity regimes (Connor et al., 2004) but at lower abundance than in this biotope (FaAlCr.Car).

Sensitivity review. CR.MCR.EcCr.FaAlCr.Car is recorded exclusively in full marine conditions (30-40 ‰) (Connor et al., 2004).  Spirobranchus triqueter is likely to be able to tolerate reduced salinity. Records from the MNCR suggest that Caryophyllia smithii, Alcyonium digitatum and Echinus esculentus can occur in reduced salinity habitats. However, their abundance is likely to decrease is the salinity decreased to 'reduced' conditions.  Therefore, resistance has been assessed as ‘Medium’, resilience as ‘High’ and sensitivity has been assessed as ‘Low’.

Water flow (tidal current) changes (local)

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

Evidence

CR.MCR.EcCr.FaAlCr.Car is recorded from weak to very weak (<0.5 m/sec) tidal streams and, although it also occurs in wave exposed to sheltered conditions it is characterized by very little water movement (Connor et al., 2004). Caryophyllia smithii is recorded from biotopes in moderately strong to very weak tidal streams. Alcyonium digitatum and Spirobranchus triqueter are suspension feeders relying on water currents to supply food. These taxa, therefore, thrive in conditions of vigorous water flow e.g. around Orkney and St Abbs, Scotland, where the community may experience tidal currents of 3 and 4 knots during spring tides (De Kluijver, 1993).Spirobranchus triqueter has been recorded in areas with very sheltered to exposed water flow rates (Price et al., 1980). Wood (1988) observed Spirobranchus sp. in strong tidal streams and Hiscock (1983) found that in strong tidal streams or strong wave action where abrasion occurs, fast growing species such as Spirobranchus triqueter occur. Echinus esculentus occurred in kelp beds on the west coast of Scotland in currents of about 0.5 m/sec. Outside the beds specimens were occasionally seen being rolled by the current (Comely & Ansell, 1988), which may have been up to 1.4 m/sec. Urchins are removed from the stipe of kelps by wave and current action. Echinus esculentus are also displaced by storm action. After disturbance Echinus esculentus migrates up the shore, an adaptation to being washed to deeper water by wave action (Lewis & Nichols, 1979a). Therefore, increased water flow may remove the population from the affected area, probably to deeper water, although individuals would probably not be killed in the process and could recolonize the area quickly.

Sensitivity assessment. CR.MCR.EcCr.FaAlCr.Car is recorded from weak to very weak (<0.5 m/sec) tidal streams and is characterized by very little water movement. A further decrease in tidal streams is 'not relevant'. However, an increase in flow of 0.1-0.2 m/s is also not likely to have a significant effect on the biological community as the important charactersitic species also occur in similar FaAlCr biotopes in stronger flow. Echinus esculentus may become dislodged but are unlikely to be killed and may recolonize quickly. Therefore, resistance has been assessed as ‘High’, resilience as ‘High’ and sensitivity has been assessed as ‘Not sensitive’ at the benchmark level.

Emergence regime changes

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

Evidence

Changes in emergence is Not relevant  fully subtidal/circalittoral biotopes. The pressure benchmark is relevant only to littoral and shallow sublittoral fringe biotopes.

Wave exposure changes (local)

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

Evidence

CR.MCR.EcCr.FaAlCr.Car is recorded from wave exposed to sheltered sites (Connor et al., 2004). Alcyonium digitatum,  Spirobranchus triqueter are suspension feeders relying on water currents to supply food. These taxa, therefore, thrive in conditions of vigorous water flow.The depth at which this biotope is recorded (10-30 m)may therefore also negate the direct physical effects of a localised change in wave height; as wave attenuation is directly related to water depth (Hiscock, 1983).

Echinus esculentus occurred in kelp beds on the west coast of Scotland in currents of about 0.5 m/sec. Outside the beds specimens were occasionally seen being rolled by the current (Comely & Ansell, 1988), which may have been up to 1.4 m/sec. Urchins are removed from the stipe of kelps by wave and current action. Echinus esculentus are also displaced by storm action. After disturbance Echinus esculentus migrates up the shore, an adaptation to being washed to deeper water by wave action (Lewis & Nichols, 1979a). Keith Hiscock (pers. comm.) reported Echinus esculentus occurred in significant numbers as shallow as 15 m below low water at the extremely wave exposed site of Rockall, Scotland. Therefore, localised increases in wave height may remove the population from the affected area, probably to deeper water, although individuals would probably not be killed in the process and could recolonize the area quickly.

Sensitivity assessment. Wave action is a fundamental environmental variable controlling the biological community of sublittoral biotopes. A large and significant change in wave height may fundamentally alter the character of FaAlCr biotopes. However, a change in near shore significant wave height of 3-5% is not likely to have a significant effect on the biological community. Resistance has been assessed as ‘High’, resilience has been assessed as ‘High’. Sensitivity has been assessed as ‘Not sensitive’.

Transition elements & organo-metal contamination

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

Evidence

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

No information on the direct biological effects of heavy metal contamination on Alcyonium digitatum. Possible sub-lethal effects of exposure to heavy metals may result in a change in morphology, growth rate or disruption of the reproductive cycle. The vulnerability of this species to concentrations of pollutants may also depend on variations in other factors e.g. temperature and salinity conditions outside the normal range. Based on the available evidence for several species Bryan (1984) suggested that polychaetes are fairly resistant to heavy metals. Bryozoans are common members of the fouling community and amongst those organisms most resistant to antifouling measures, such as copper containing anti-fouling paints (Soule & Soule, 1979; Holt et al., 1995). Bryozoans were shown to bio accumulate heavy metals to a certain extent (Holt et al., 1995). For example, Bowerbankia gracialis and Nolella pusilla accumulated Cd, exhibiting sublethal effects (reduced sexual reproduction and inhibited resting spore formation) between 10-100 µg Cd /l and fatality above 500 µg Cd/l (Kayser, 1990). Little is known about the effects of heavy metals on echinoderms. Bryan (1984) reported that early work had shown that echinoderm larvae were sensitive to heavy metals contamination, for example, Migliaccio et al. (2014) reported exposure of Paracentrotus lividis larvae to increased levels of cadmium and manganese caused abnormal larval development and skeletal malformations. Kinne (1984) reported developmental disturbances in Echinus esculentus exposed to waters containing 25 µg / l of copper (Cu).

Bryozoans are common members of the fouling community and amongst those organisms most resistant to antifouling measures, such as copper containing anti-fouling paints (Soule & Soule, 1979; Holt et al., 1995). Bryan & Gibbs (1991) reported that there was little evidence regarding TBT toxicity in bryozoa with the exception of the encrusting Schizoporella errata, which suffered 50% mortality when exposed for 63 days to 100 ng/l TBT. Rees et al. (2001) reported that the abundance of epifauna (including bryozoans) had increased in the Crouch estuary in the 5 years since TBT was banned from use on small vessels. This last report suggests that bryozoans may be at least inhibited by the presence of TBT.

Hydrocarbon & PAH contamination

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

Evidence

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

CR.MCR.EcCr.FaAlCr biotopes are subtidal biotopes (Connor et al., 2004). Oil pollution is mainly a surface phenomenon its impact upon circalittoral turf communities is likely to be limited. However, as in the case of the Prestige oil spill off the coast of France, high swell and winds can cause oil pollutants to mix with the seawater and potentially negatively affect sublittoral habitats (Castège et al., 2014). Smith (1968) reported dead colonies of Alcyonium digitatum at a depth of 16 m in the locality of Sennen Cove, Cornwall, which was likely a result of toxic detergents sprayed along the shoreline to disperse oil from the Torrey Canyon tanker spill (Budd, 2008). Little information on the effects of hydrocarbons on bryozoans was found. Ryland & Putron (1998) did not detect adverse effects of oil contamination on the bryozoan Alcyonidium spp. in Milford Haven or St. Catherine's Island, south Pembrokeshire although it did alter the breeding period. Large numbers of dead polychaetes and other fauna were washed up at Rulosquet Marsh near the Isle de Grand following the Amoco Cadiz oil spill in 1978 (Cross et al., 1978). However, no information was found relating to Spirobranchus triqueter in particular.

Echinus esculentus is subtidal and unlikely to be directly exposed to oil spills. However, as with the ‘Prestige’ oil spill rough seas can cause mixing with the oil and the seawater, and therefore sub-tidal habitats can be affected by the oil spill. Castège et al., (2014) recorded the recovery of rocky shore communities following the Prestige oil spill which impacted the French Atlantic coast. Rough weather at the time of the spill increased mixing between the oil and seawater, causing subtidal communities/habitats to be affected. The urchin Echinus esculentus was reported absent after the oil spill, however, returned after 2-5 years. Large numbers of dead Echinus esculentus were found between 5.5 and 14.5 m in the vicinity of Sennen cove, presumably due to a combination of wave exposure and heavy spraying of dispersants following the Torrey canyon oil spill (Smith, 1968). Smith (1968) also demonstrated that 0.5 -1 ppm of the detergent BP1002 resulted in developmental abnormalities in its echinopluteus larvae. Echinus esculentus populations in the vicinity of an oil terminal in A Coruna Bay, Spain, showed developmental abnormalities in the skeleton. The tissues contained high levels of aliphatic hydrocarbons, naphthalenes, pesticides and heavy metals (Zn, Hg, Cd, Pb, and Cu) (Gomez & Miguez-Rodriguez, 1999).

Synthetic compound contamination

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

Evidence

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

Smith (1968) reported dead colonies of Alcyonium digitatum at a depth of 16 m in the locality of Sennen Cove, Cornwall resulting from the offshore spread and toxic effect of detergents (a mixture of a surfactant and an organic solvent) e.g. BP 1002 sprayed along the shoreline to disperse oil from the Torrey Canyon tanker spill. Possible sub-lethal effects of exposure to synthetic chemicals may result in a change in morphology, growth rate or disruption of the reproductive cycle. The vulnerability of this species to concentrations of pollutants may also depend on variations in other factors e.g. temperature and salinity conditions outside the normal range (Budd, 2008).

Hoare & Hiscock (1974) suggested that polyzoa (bryozoa) were amongst the most intolerant species to acidified halogenated effluents in Amlwch Bay, Anglesey and reported that Flustra foliacea did not occur less than 165m from the effluent source. The evidence, therefore, suggests that Securiflustra securifrons would be sensitive to synthetic compounds.

Large numbers of dead Echinus esculentus were found between 5.5 and 14.5 m in the vicinity of Sennen, presumably due to a combination of wave exposure and heavy spraying of dispersants in that area following the Torrey Canyon oil spill (Smith 1968). Smith (1968) also demonstrated that 0.5 -1ppm of the detergent BP1002 resulted in developmental abnormalities in echinopluteus larvae of Echinus esculentus. Echinus esculentus populations in the vicinity of an oil terminal in A Coruna Bay, Spain, showed developmental abnormalities in the skeleton. The tissues contained high levels of aliphatic hydrocarbons, naphthalenes, pesticides and heavy metals (Zn, Hg, Cd, Pb, and Cu) (Gomez & Miguez-Rodriguez 1999).

Radionuclide contamination

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

Evidence

No evidence was found

Introduction of other substances

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

Evidence

This pressure is Not assessed.

De-oxygenation

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

Evidence

In general, respiration in most marine invertebrates do not appear to be significantly affected until extremely low concentrations are reached. For many benthic invertebrates, this concentration is about 2 ml/l or even less (Herreid, 1980; Rosenberg et al., 1991; Diaz & Rosenberg, 1995).

Alcyonium digitatum mainly inhabits environments in which the oxygen concentration usually exceeds 5 ml/l and respiration is aerobic (Budd, 2008). In August 1978 a dense bloom of a dinoflagellate, Gyrodinium aureolum occurred surrounding Geer reef in Penzance Bay, Cornwall and persisted until September that year. Observations by local divers indicated a decrease in underwater visibility (<1 m) from below 8 m Below Sea Level. It was also noted that many of the faunal species appeared to be affected, e.g. no live Echinus esculentus were observed whereas on surveys prior to August were abundant, Alcyonium sp. and Bryozoans were also in an impoverished state. Caryophyllia smithii were also in a contracted state, apparently dead, and with Echinus esculentus were the worst affected species during the bloom.  During follow up surveys conducted in early September Alcyonium sp. were noted to be much healthier and feeding. It was suggested the decay of Gyrodinium aureolum either reduced oxygen levels or physically clogged faunal feeding mechanisms. Adjacent reefs where also surveyed during the same time period and the effects of the Gyrodinium aureolum bloom were less apparent. It was suggested that higher water agitation in shallow water on reefs more exposed to wave action were less affected by the phytoplankton bloom (Griffiths et al., 1979).

Sensitivity assessment. There is anecdotal evidence to suggest that Alcyonium digitatum, Caryophyllia smithii and Echinus esculentus are sensitive to hypoxic events.  Loss of the important characteristic species and the major grazer could result in a change in the biotope. Therefore, a resistance of Low is suggested. However, because the degree of deoxygenation wasn’t quantified the evidence cannot be compared to the pressure benchmark and confidence in the assessment is 'Low'. Resilience is probably High so that sensitivity is assessed as Low.

Nutrient enrichment

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

Evidence

This biotope is considered to be 'Not sensitive' at the pressure benchmark that assumes compliance with good status as defined by the WFD.

Alcyonium digitatum and Caryophyllia smithii are suspension feeders of phytoplankton and zooplankton. Nutrient enrichment of coastal waters that enhances the population of phytoplankton may be beneficial to Alcyonium digitatum and Caryophyllia smithii in terms of an increased food supply but the effects are uncertain (Hartnoll, 1998). The survival of Alcyonium digitatum and Caryophyllia smithii may be influenced indirectly. High primary productivity in the water column combined with high summer temperature and the development of thermal stratification (which prevents mixing of the water column) can lead to hypoxia of the bottom waters which faunal species are likely to be highly intolerant of (see de-oxygenation pressure).

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types.

It was suggested by Comely & Ansell (1988) that Echinus esculentus could absorb dissolved organic material for the purposes of nutrition. Nutrient enrichment may encourage the growth of ephemeral and epiphytic algae and therefore increase sea-urchin food availability. Lawrence (1975) reported that sea urchins had persisted over 13 years on barren grounds near sewage outfalls, presumably feeding on dissolved organic material, detritus, plankton and microalgae, although individuals died at an early age. 

Organic enrichment

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

Evidence

Alcyonium digitatum and Caryophyllia smithii are suspension feeders of phytoplankton and zooplankton. Organic enrichment of coastal waters that enhances the population of phytoplankton may be beneficial to Alcyonium digitatum and Caryophyllia smithii in terms of an increased food supply but the effects are uncertain (Hartnoll, 1998). The survival of Alcyonium digitatum and Caryophyllia smithii may be influenced indirectly. High primary productivity in the water column combined with high summer temperature and the development of thermal stratification (which prevents mixing of the water column) can lead to hypoxia of the bottom waters which faunal species are likely to be highly intolerant of (see de-oxygenation pressure).

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types.

It was suggested by Comely & Ansell (1988) that Echinus esculentus could absorb dissolved organic material for the purposes of nutrition. Organic enrichment may encourage the growth of ephemeral and epiphytic algae and therefore increase sea-urchin food availability. Lawrence (1975) reported that sea urchins had persisted over 13 years on barren grounds near sewage outfalls, presumably feeding on dissolved organic material, detritus, plankton and microalgae, although individuals died at an early age.

Sensitivity assessment. Organic enrichment is not likely to directly negatively affect the characterizing species within this biotope, however, chronic organic enrichment may cause secondary effects such as hypoxia. Resistance has been assessed as ‘Medium’, resilience as ‘High’, and sensitivity has been assessed as ‘Low’.

Physical loss (to land or freshwater habitat)

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

Evidence

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

Physical change (to another seabed type)

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

Evidence

If rock were replaced with sediment, this would represent a fundamental change to the physical character of the biotope and the species would be unlikely to recover. The biotope would be lost.

Sensitivity assessment. Resistance to the pressure is considered ‘None’, and resilience ‘Very low’. Sensitivity has been assessed as ‘High’.

Physical change (to another sediment type)

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

Evidence

Not relevant

Habitat structure changes - removal of substratum (extraction)

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

Evidence

The species characterizing this biotope are epifauna or epiflora occurring on rock and would be sensitive to the removal of the habitat. However, extraction of rock substratum is considered unlikely and this pressure is considered to be ‘Not relevant’ to hard substratum habitats.

Abrasion / disturbance of the surface of the substratum or seabed

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

Evidence

CR.MCR.EcCr.FaAlCr biotopes are subtidal habitats (Connor et al., 2004). Therefore, abrasion is most likely to be a result of bottom or pot fishing gear, cable laying etc., which may cause localised mobility of the substrata and mortality of the resident community. The effect would be situation dependent, however, if bottom fishing gear were towed over a site it may mobilise a high proportion of the rock substrata and cause high mortality in the resident community.

Alcyonium digitatum, Echinus esculentus, and Caryophyllia smithii are sessile or slow moving species that might be expected to suffer from the effects of dredging. Boulcott & Howell (2011) conducted experimental Newhaven scallop dredging over a circalittoral rock habitat in the sound of Jura, Scotland and recorded the damage to the resident community. The results indicated that the sponge Pachymatisma johnstoni was highly damaged by the experimental trawl. However, only 13% of photographic samples showed visible damage to Alcyonium digitatum. Where Alcyonium digitatum damage was evident it tended to be small colonies that were ripped off the rock. The authors highlight physical damage to faunal turfs (erect bryozoans and hydroids) was difficult to quantify in the study. However, the faunal turf communities did not show large signs of damage and were only damaged by the scallop dredge teeth which was often limited in extent (approximately 2 cm wide tracts). The authors indicated that species such as Alcyonium digitatum and faunal turf communities were not as vulnerable to damage through trawling as sedimentary fauna and whilst damage to circalittoral rock fauna did occur it was of an incremental nature, with loss of species such as Alcyonium digitatum and faunal turf communities increasing with repeated trawls.

Species with fragile tests, such as Echinus esculentus were reported to suffer badly as a result of scallop or queen scallop dredging (Bradshaw et al., 2000; Hall-Spencer & Moore, 2000a). Kaiser et al. (2000) reported that Echinus esculentus were less abundant in areas subject to high trawling disturbance in the Irish Sea. Jenkins et al. (2001) conducted experimental scallop trawling in the North Irish sea and recorded the damage caused to several conspicuous megafauna species, both when caught as by-catch and when left on the seabed. The authors predicted 16.4% of Echinus esculentus were crushed/dead, 29.3% would have >50% spine loss/minor cracks, 1.1% would have <50% spine loss and the remaining 53.3% would be in good condition. Sea urchins can rapidly regenerate spines, e.g. Psammechinus miliaris were found to re-grow all spines within a period of 2 months (Hobson, 1930).  The trawling examples mentioned above were conducted on sedimentary habitats and thus the evidence is not directly relevant to the rock based biotopes, however, does indicate the likely effects of abrasion on Echinus esculentus.

Sensitivity assessment. Resistance has been assessed ‘Medium’, resilience has been assessed as ‘High’. Sensitivity has been assessed as ‘Low’. Please note, Boulcott & Howell (2011) did not mention the abrasion caused by fully loaded collection bags on the new haven dredges. A fully loaded Newhaven dredge may cause higher damage to the community as indicated in their study.

Penetration or disturbance of the substratum subsurface

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

Evidence

The species characterizing this biotope group are epifauna or epiflora occurring on rock which is resistant to subsurface penetration.  The assessment for abrasion at the surface only is therefore considered to equally represent sensitivity to this pressure. This pressure is not thought relevant to hard rock biotopes.

Changes in suspended solids (water clarity)

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

Evidence

Alcyonium digitatum and Caryophyllia smithii are not thought highly susceptible to changes in water clarity due to the fact they are suspension feeding organisms and are not directly dependent on sunlight for nutrition. Alcyonium digitatum has been shown to be tolerant of high levels of suspended sediment. Hill et al. (1997) demonstrated that Alcyonium digitatum sloughed off settled particles with a large amount of mucous. Alcyonium digitatum is also known to inhabit the entrances to sea lochs (Budd, 2008) or the entrances to estuaries (Braber & Borghouts, 1977) where water clarity is likely to be highly variable.

Moore (1977a) suggested that Echinus esculentus was unaffected by turbid conditions. Echinus esculentus is an important grazer of red macro-algae within CR.MCR.EcCr. Increased turbidity and resultant reduced light penetration is likely to negatively affect algal growth. However, Echinus esculentus can feed on alternative prey, detritus or dissolved organic material (Lawrence, 1975, Comely & Ansell, 1988).

According to Bacescu (1972), sabellids are accustomed to turbidity and silt. Spirobranchus triqueter has also recently been recorded by De Kluijver (1993) from Scotland in the aphotic zone, indicating that the species would not be sensitive to light reduction due to an increase in turbidity. In addition, Spirobranchus triqueter is commonly found under boulders where sediment may accumulate.

Sensitivity assessment. Resistance has been assessed as ‘High’, resilience as ‘High’ and sensitivity has been assessed as ‘Not Sensitive’.

Smothering and siltation rate changes (light)

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

Evidence

Alcyonium digitatum and Caryophyllia smithii are sessile and thus would be unable to avoid the deposition of a smothering layer of sediment. Some Alcyonium digitatum colonies can attain a height of up to 20 cm (Budd, 2008), while Echinus esculentus are large globular urchins which can reach a diameter of 17 cm (Tyler-Walters, 2008). Spirobranchus triqueter are an encrusting species and would thus likely be smothered, and depending on sediment retention could block larval settlement.

Caryophyllia smithii is small (approx. <3 cm height from the seabed) and would therefore likely be inundated in a “light” sedimentation event. However, Bell & Turner (2000) reported that Caryophyllia smithii was abundant at sites of “moderate” sedimentation (7mm ± 0.5mm) in Lough Hyne. It is, therefore, likely that Caryophyllia smithii would be resistant to periodic sedimentation. If 5 cm of sediment were removed rapidly, via tidal currents, Caryophyllia smithii would likely remain within the biotope.

Comely & Ansell (1988) recorded large Echinus esculentus from kelp beds on the west coast of Scotland, in which, the substratum was seasonally covered with "high levels" of silt. This suggests that Echinus esculentus is unlikely to be killed by smothering, however, smaller specimens and juveniles may be less resistant. A layer of sediment may interfere with larval settlement.  If retained within the host biotope for extended periods a layer of 5 cm of the sediment may negatively affect successive recruitment events.

Sensitivity assessment. CR.MCR.EcCr.FaAlCr.Car is recorded from areas with very little water movement (Connor et al., 2004) so that any deposit of sediment is likely to remain for some time and only vertical surfaces escape smothering.  Echinus may be deterred from feeding but small encrusting forms (Caryophyllia smithii and Spirobranchus triqueter) may be smothered and prevented from feeding or exposed to anoxia. Resistance has been assessed as ‘Low’, resilience as ‘High’, and sensitivity has, therefore, been assessed as ‘Low’.

Smothering and siltation rate changes (heavy)

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

Evidence

Alcyonium digitatum and Caryophyllia smithii are sessile and thus would be unable to avoid the deposition of a smothering layer of sediment. Some Alcyonium digitatum colonies can attain a height of up to 20 cm (Budd, 2008), while Echinus esculentus are large globular urchins which can reach a diameter of 17 cm (Tyler-Walters, 2008). Spirobranchus triqueter are an encrusting species and would thus likely be smothered, and depending on sediment retention could block larval settlement.

Caryophyllia smithii is small (approx. <3 cm height from the seabed) and would therefore likely be inundated in a “light” sedimentation event. However, Bell & Turner (2000) reported that Caryophyllia smithii was abundant at sites of “moderate” sedimentation (7mm ± 0.5mm) in Lough Hyne. It is, therefore, likely that Caryophyllia smithii would be resistant to periodic sedimentation. If 5 cm of sediment were removed rapidly, via tidal currents, Caryophyllia smithii would likely remain within the biotope.

Comely & Ansell (1988) recorded large Echinus esculentus from kelp beds on the west coast of Scotland, in which, the substratum was seasonally covered with "high levels" of silt. This suggests that Echinus esculentus is unlikely to be killed by smothering, however, smaller specimens and juveniles may be less resistant. A layer of sediment may interfere with larval settlement.  If retained within the host biotope for extended periods a layer of 5 cm of the sediment may negatively affect successive recruitment events.

Sensitivity assessment. CR.MCR.EcCr.FaAlCr.Car is recorded from areas with very little water movement (Connor et al., 2004) so that any deposit of sediment is likely to remain for some time and only vertical surfaces escape smothering.  Echinus may be deterred from feeding but small encrusting forms (Caryophyllia smithii and Spirobranchus triqueter) may be smothered and prevented from feeding or exposed to anoxia. Resistance has been assessed as ‘Low’, resilience as ‘High’, and sensitivity has, therefore, been assessed as ‘Low’.

Litter

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

Evidence

Not assessed.

Electromagnetic changes

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

Evidence

There was no evidence on which to assess this pressure. 

Underwater noise changes

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

Evidence

Alcyonium digitatum, Echinus esculentus and Caryophyllia smithii have no hearing perception but vibrations may cause an impact, however no studies exist to support an assessment.

Introduction of light or shading

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

Evidence

There is no evidence to suggest that If exposed to anthropogenic light sources algal species would benefit. CR.MCR.EcCr.FaAlCr biotopes are circalittoral and are thus by definition naturally shaded environments with low light levels. Increased shading (e.g. by the construction of a pontoon, pier etc) could be beneficial to the characterizing species within these biotopes.

Sensitivity assessment. Resistance is probably 'High', with a 'High' resilience and a sensitivity of 'Not Sensitive'. 

Barrier to species movement

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

Evidence

Not relevant: barriers and changes in tidal excursion are not relevant to biotopes restricted to open waters.

Death or injury by collision

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

Evidence

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

Visual disturbance

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

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 that may result in changes in the genetic structure of local populations, hybridization, or change in community structure. Further detail

Evidence

Alcyonium digitatum, Securiflustra securifrons & Spirobranchus triqueter are not cultivated or translocated. Echinus esculentus was identified by Kelly & Pantazis (2001) as a species suitable for culture for the urchin Roe industry. However, no evidence could be found to suggest that significant Echinus esculentus mariculture was present in the UK. If industrially cultivated it is feasible that Echinus esculentus individuals could be translocated.

Translocation also has the potential to transport pathogens to uninfected areas (see pressure ‘introduction of microbial pathogens’). The sensitivity of the ‘donor’ population to harvesting to supply stock for translocation is assessed for the pressure ‘removal of target species’.

Introduction or spread of invasive non-indigenous species

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

Evidence

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

Didemnum vexillum is an invasive colonial sea squirt native to Asia which was first recorded in the UK in Darthaven Marina, Dartmouth in 2005. Didemnum vexillum can form extensive mats over the substrata it colonizes; binding boulders and cobbles and altering the host habitat (Griffith et al., 2009). Didemnum vexillum can also grow over and smother the resident biological community. Recent surveys within Holyhead Marina, North Wales have found Didemnum vexillum growing on and smothering native tunicate communities (Griffith et al., 2009). Due to the rapid-re-colonization of Didemnum vexillum eradication attempts have to date failed. 

Presently Didemnum vexillum is isolated to several sheltered locations in the UK (NBN, 2015), however, Didemnum vexillum has successfully colonized the offshore location of the Georges Bank, USA (Lengyel et al., 2009) which is more exposed than the locations that Didemnum vexillum has colonized in the UK. It is, therefore, possible that Didemnum vexillum could colonize more exposed locations within the UK and could, therefore, pose a threat to CR.MCR.EcCr.FaAlCr biotopes in the future.

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

Introduction of microbial pathogens

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

Evidence

No evidence was found to suggest that any of the characterizing species within CR.MCR.EcCr.FaAlCr biotopes are sensitive to current/known microbial pathogens.

Alcyonium digitatum acts as the host for the endoparasitic species Enalcyonium forbesiand and Enalcyonium rubicundum (Stock, 1988). Parasitisation may reduce the viability of a colony but not to the extent of killing them but no further evidence was found to substantiate this suggestion. Thomas (1940) recorded parasites of Spirobranchus triqueter. Trichodina pediculus (a ciliate) was observed in high numbers moving over the branchial crown. However, this relationship is symbiotic, not parasitic. Parasites found in the worm include gregarines & ciliated protozoa and parasites that had the appearance of sporozoan cysts. However, no information was found about the effects of microbial pathogens on Spirobranchus triqueter.

Echinus esculentus is susceptible to 'Bald-sea-urchin disease', which causes lesions, loss of spines, tube feet, pedicellariae, destruction of the upper layer of skeletal tissue and death. It is thought to be caused by the bacteria Vibrio anguillarum and Aeromonas salmonicida. Bald sea-urchin disease was recorded from Echinus esculentus on the Brittany Coast. Although associated with mass mortalities of Strongylocentrotus franciscanus in California and Paracentrotus lividus in the French Mediterranean it is not known if the disease induces mass mortality (Bower, 1996).

Removal of target species

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

Evidence

None of the characterizing species within CR.MCR.EcCr.FaAlCr biotopes are commercially exploited. This pressure is considered ‘Not Relevant’.

Removal of non-target species

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

Evidence

Alcyonium digitatum and faunal turf communities  are probably resistant to abrasion through bottom fishing (see abrasion pressure). Alcyonium digitatum goes through an annual cycle, From February to July all Alcyonium digitatum colonies are feeding, from July to November, an increasing number of colonies stop feeding. During this period, a large number of polyps can retract and a variety of filamentous algae, hydroids and amphipods can colonize the surface of colonies epiphytically. From December to February, the epiphytic community is however sloughed off (Hartnoll, 1975). If Alcyonium digitatum were removed the epiphytic species would likely colonize rock surfaces and are therefore not dependent on Alcyonium digitatum.

Alcyonium digitatum and Caryophylia smithii, probably compete for space, however, there isn’t any evidence to suggest other interspecific relationships or dependencies between these species. Therefore, removal of one or a number of these species would provide colonization space and most likely benefit the species with rapid colonization rates (e.g. Spirobranchus triqueter). Echinus esculentus is an important red algal grazer  (Connor et al., 2004), without which the abundance of red algae may increase and possibly displace some of the faunal turf species.

Sensitivity assessment. If Alcyonium digitatum, Caryophylia smithii or Echinus were removed (e.g. as incidental by-catch) the character of the biotope would probably change. Therefore, resistance has been assessed as ‘Low’, resilience has been assessed as ’High’. Sensitivity has been assessed as ‘Low’.