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

The distribution of seaweeds is climatically defined (Kain, 1979, Van Den Hoek, 1982, Breeman, 1990, Lüning, 1990). Northern boundaries are set by winter temperatures that are lethal or summer temperatures too low for growth and/or reproduction, whilst southern limits are set by high lethal summer temperatures or winter temperatures too high for induction of a crucial step in the life cycle (Breeman, 1990). Kelps have a high dependence on ocean temperatures, which makes them highly vulnerable to ocean warming (Assis et al., 2014). As temperatures increase, populations found towards the upper limit of their temperature range may be adversely affected by warming as physiological thresholds are exceeded (Wiens, 2016). Thermal stress can lead to mortality and consequent population-level effects, such as decreased abundance, altered size structure, local extinction and range contractions (Smale, 2019).

Alaria esculenta is a polar to cold-temperate species that has been recorded from Brittany, France to Northern Norway (Birkett et al., 1998b). Sea temperature regulates metabolism and reproduction and defines the regional distribution of Alaria esculenta (Fredersdorf et al., 2009). The southern limit of Alaria esculenta has been defined at the 20°C isotherm (Munda & Lüning, 1977; Fredersdorf et al., 2009), however, it is more common north of the 16°C isotherm (Munda & Lüning, 1977; Van der Hoek, 1982). As a result of this upper temperature threshold, Alaria esculenta is largely absent from the southern North Sea and English channel where summer temperatures exceed 16°C and has already shown signs of contracting its range in the UK as a result of ocean warming (Mieszkowska, 2016).

Munda & Lüning (1977) observed that temperatures of 16-17°C, sustained over 2 weeks in Helgoland, Germany, were lethal to resident Alaria esculenta. Experimental observations showed that acute exposure to ≥21°C is lethal to Alaria esculenta causing bleaching and disintegration (Sundene, 1962; Fredersdorf et al., 2009). At its northern range edge (Svalbard), it is a prominent macroalga on sublittoral fringe bedrock.  At these northern latitudes, average summer temperature can reach 5°C, with an average annual sea temperature of 3°C (1980-2014, Beszczynska-Möller & Dye, 2013). Experimental observations conducted by Fredersdorf et al. (2009) found the optimal temperature for sporophyte photosynthesis was within the range of 13-17°C, however, the optimal temperature for Alaria esculenta germination was 2-12°C (Fredersdorf et al., 2009).

Alaria esculenta is nearing its southern limits in the UK, with this species only occurring as far south as Brittany, France. Sea surface temperatures (SST) around the UK currently fall between 6-19°C (Huthnance, 2010). The available evidence suggests that the effects of ocean warming may occur throughout the year.  If winter temperatures exceed 12°C this is likely to inhibit germination, whilst summer temperatures greater than 20°C will cause mortality of Alaria esculenta.

Laminaria digitata is a polar to cold-temperate Atlantic species, which grows at temperatures of between 0-20 °C, with optimum growth at 10 °C (Bolton & Lüning, 1982), although both growth and photosynthesis show a significant decline at 18°C (Hargrave et al., 2017). Under experimental conditions, Laminaria digitata did not survive during 9 weeks of exposure to temperatures of 23°C (Wilson et al., 2015). During an exceptionally warm summer in Norway, Sundene (1964) reported mortality of sporophytes exposed to temperatures of 22-23°C.

Corallina officinalis may tolerate between -4 to 28°C (Lüning, 1990), although when Colthart & Johansen (1973) exposed this species to a number of different temperatures, they found that growth was maintained at 18°C and ceased at 25°C. Abrupt temperature changes (10°C in California, Seapy & Littler 1984; 4.8 to 8.5°C, Hawkins & Hartnoll, 1985) resulted in dramatic declines. However, in both cases recovery was rapid, suggesting that the crustose bases survived.

Sensitivity assessment. Alaria esculenta is competitively inferior to Laminaria digitata in the northeast Atlantic (Hawkins & Harkin, 1985) where it is confined to more exposed shores and can colonize sheltered environments only if Laminaria digitata is removed. As temperatures rise to up to 20°C, it is expected that the competitive ability of this species will decrease further, while as temperatures rise above 20°C both Alaria esculenta and Laminaria digitata will be lost, with more, warm adapted, algae taking their place. Brodie et al. (2014) predict that Alaria esculenta will be completely lost from Boreal habitats by the end of this century, whilst all kelp species will be lost from the Lusitanian region, replaced by smaller, fleshy algae.

Sea surface temperatures around the UK are currently between 6-19°C (Huthnance, 2010). Under the middle emission scenario, Alaria esculenta is likely to be lost from most of the UK, although it may manage to maintain some populations in Scotland, where current summer temperatures often reach 14°C, leading to potential summer temperatures of 17°C. Laminaria digitata is likely to be lost in southern parts of the UK, where summer temperatures will exceed 20°C, but will likely remain present around the rest of the UK. Corallina officinalis and other warm-tolerant species should be able to cope with temperature increases under a middle emission scenario. As Alaria esculenta is the main characterizing species for this study and is expected to be lost from a large portion of the UK, under this scenario resistance is assessed as ‘Low’. Resilience is assessed as ‘Very Low’, as the loss is likely to be a long-term decline, due to the long-term nature of ocean warming. Therefore, this biotope is assessed as having ‘High’ sensitivity to ocean warming under a middle emission scenario.

For the high emission scenario and extreme scenario, sea temperatures may rise by 4-5°C to give potential southern summer temperatures of 23-24°C and northern summer temperatures of 18-19°C. Under these scenarios, it is likely that Alaria esculenta will be lost almost completely from the UK, and Laminaria digitata will retreat northwards, with large losses across England, Ireland, and Wales. Therefore, resistance is assessed as ‘None’, and resilience is assessed as ‘Very low’. Overall, this biotope is assessed as having ‘High’ sensitivity to ocean warming for the high emission scenario and the extreme scenario.  

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

The distribution of seaweeds is climatically defined (Kain, 1979, Van Den Hoek, 1982, Breeman, 1990, Lüning, 1990). Northern boundaries are set by winter temperatures that are lethal or summer temperatures too low for growth and/or reproduction, whilst southern limits are set by high lethal summer temperatures or winter temperatures too high for induction of a crucial step in the life cycle (Breeman, 1990). Kelps have a high dependence on ocean temperatures, which makes them highly vulnerable to ocean warming (Assis et al., 2014). As temperatures increase, populations found towards the upper limit of their temperature range may be adversely affected by warming as physiological thresholds are exceeded (Wiens, 2016). Thermal stress can lead to mortality and consequent population-level effects, such as decreased abundance, altered size structure, local extinction and range contractions (Smale, 2019).

Alaria esculenta is a polar to cold-temperate species that has been recorded from Brittany, France to Northern Norway (Birkett et al., 1998b). Sea temperature regulates metabolism and reproduction and defines the regional distribution of Alaria esculenta (Fredersdorf et al., 2009). The southern limit of Alaria esculenta has been defined at the 20°C isotherm (Munda & Lüning, 1977; Fredersdorf et al., 2009), however, it is more common north of the 16°C isotherm (Munda & Lüning, 1977; Van der Hoek, 1982). As a result of this upper temperature threshold, Alaria esculenta is largely absent from the southern North Sea and English channel where summer temperatures exceed 16°C and has already shown signs of contracting its range in the UK as a result of ocean warming (Mieszkowska, 2016).

Munda & Lüning (1977) observed that temperatures of 16-17°C, sustained over 2 weeks in Helgoland, Germany, were lethal to resident Alaria esculenta. Experimental observations showed that acute exposure to ≥21°C is lethal to Alaria esculenta causing bleaching and disintegration (Sundene, 1962; Fredersdorf et al., 2009). At its northern range edge (Svalbard), it is a prominent macroalga on sublittoral fringe bedrock.  At these northern latitudes, average summer temperature can reach 5°C, with an average annual sea temperature of 3°C (1980-2014, Beszczynska-Möller & Dye, 2013). Experimental observations conducted by Fredersdorf et al. (2009) found the optimal temperature for sporophyte photosynthesis was within the range of 13-17°C, however, the optimal temperature for Alaria esculenta germination was 2-12°C (Fredersdorf et al., 2009).

Alaria esculenta is nearing its southern limits in the UK, with this species only occurring as far south as Brittany, France. Sea surface temperatures (SST) around the UK currently fall between 6-19°C (Huthnance, 2010). The available evidence suggests that the effects of ocean warming may occur throughout the year.  If winter temperatures exceed 12°C this is likely to inhibit germination, whilst summer temperatures greater than 20°C will cause mortality of Alaria esculenta.

Laminaria digitata is a polar to cold-temperate Atlantic species, which grows at temperatures of between 0-20 °C, with optimum growth at 10 °C (Bolton & Lüning, 1982), although both growth and photosynthesis show a significant decline at 18°C (Hargrave et al., 2017). Under experimental conditions, Laminaria digitata did not survive during 9 weeks of exposure to temperatures of 23°C (Wilson et al., 2015). During an exceptionally warm summer in Norway, Sundene (1964) reported mortality of sporophytes exposed to temperatures of 22-23°C.

Corallina officinalis may tolerate between -4 to 28°C (Lüning, 1990), although when Colthart & Johansen (1973) exposed this species to a number of different temperatures, they found that growth was maintained at 18°C and ceased at 25°C. Abrupt temperature changes (10°C in California, Seapy & Littler 1984; 4.8 to 8.5°C, Hawkins & Hartnoll, 1985) resulted in dramatic declines. However, in both cases recovery was rapid, suggesting that the crustose bases survived.

Sensitivity assessment. Alaria esculenta is competitively inferior to Laminaria digitata in the northeast Atlantic (Hawkins & Harkin, 1985) where it is confined to more exposed shores and can colonize sheltered environments only if Laminaria digitata is removed. As temperatures rise to up to 20°C, it is expected that the competitive ability of this species will decrease further, while as temperatures rise above 20°C both Alaria esculenta and Laminaria digitata will be lost, with more, warm adapted, algae taking their place. Brodie et al. (2014) predict that Alaria esculenta will be completely lost from Boreal habitats by the end of this century, whilst all kelp species will be lost from the Lusitanian region, replaced by smaller, fleshy algae.

Sea surface temperatures around the UK are currently between 6-19°C (Huthnance, 2010). Under the middle emission scenario, Alaria esculenta is likely to be lost from most of the UK, although it may manage to maintain some populations in Scotland, where current summer temperatures often reach 14°C, leading to potential summer temperatures of 17°C. Laminaria digitata is likely to be lost in southern parts of the UK, where summer temperatures will exceed 20°C, but will likely remain present around the rest of the UK. Corallina officinalis and other warm-tolerant species should be able to cope with temperature increases under a middle emission scenario. As Alaria esculenta is the main characterizing species for this study and is expected to be lost from a large portion of the UK, under this scenario resistance is assessed as ‘Low’. Resilience is assessed as ‘Very Low’, as the loss is likely to be a long-term decline, due to the long-term nature of ocean warming. Therefore, this biotope is assessed as having ‘High’ sensitivity to ocean warming under a middle emission scenario.

For the high emission scenario and extreme scenario, sea temperatures may rise by 4-5°C to give potential southern summer temperatures of 23-24°C and northern summer temperatures of 18-19°C. Under these scenarios, it is likely that Alaria esculenta will be lost almost completely from the UK, and Laminaria digitata will retreat northwards, with large losses across England, Ireland, and Wales. Therefore, resistance is assessed as ‘None’, and resilience is assessed as ‘Very low’. Overall, this biotope is assessed as having ‘High’ sensitivity to ocean warming for the high emission scenario and the extreme scenario.  

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

The distribution of seaweeds is climatically defined (Kain, 1979, Van Den Hoek, 1982, Breeman, 1990, Lüning, 1990). Northern boundaries are set by winter temperatures that are lethal or summer temperatures too low for growth and/or reproduction, whilst southern limits are set by high lethal summer temperatures or winter temperatures too high for induction of a crucial step in the life cycle (Breeman, 1990). Kelps have a high dependence on ocean temperatures, which makes them highly vulnerable to ocean warming (Assis et al., 2014). As temperatures increase, populations found towards the upper limit of their temperature range may be adversely affected by warming as physiological thresholds are exceeded (Wiens, 2016). Thermal stress can lead to mortality and consequent population-level effects, such as decreased abundance, altered size structure, local extinction and range contractions (Smale, 2019).

Alaria esculenta is a polar to cold-temperate species that has been recorded from Brittany, France to Northern Norway (Birkett et al., 1998b). Sea temperature regulates metabolism and reproduction and defines the regional distribution of Alaria esculenta (Fredersdorf et al., 2009). The southern limit of Alaria esculenta has been defined at the 20°C isotherm (Munda & Lüning, 1977; Fredersdorf et al., 2009), however, it is more common north of the 16°C isotherm (Munda & Lüning, 1977; Van der Hoek, 1982). As a result of this upper temperature threshold, Alaria esculenta is largely absent from the southern North Sea and English channel where summer temperatures exceed 16°C and has already shown signs of contracting its range in the UK as a result of ocean warming (Mieszkowska, 2016).

Munda & Lüning (1977) observed that temperatures of 16-17°C, sustained over 2 weeks in Helgoland, Germany, were lethal to resident Alaria esculenta. Experimental observations showed that acute exposure to ≥21°C is lethal to Alaria esculenta causing bleaching and disintegration (Sundene, 1962; Fredersdorf et al., 2009). At its northern range edge (Svalbard), it is a prominent macroalga on sublittoral fringe bedrock.  At these northern latitudes, average summer temperature can reach 5°C, with an average annual sea temperature of 3°C (1980-2014, Beszczynska-Möller & Dye, 2013). Experimental observations conducted by Fredersdorf et al. (2009) found the optimal temperature for sporophyte photosynthesis was within the range of 13-17°C, however, the optimal temperature for Alaria esculenta germination was 2-12°C (Fredersdorf et al., 2009).

Alaria esculenta is nearing its southern limits in the UK, with this species only occurring as far south as Brittany, France. Sea surface temperatures (SST) around the UK currently fall between 6-19°C (Huthnance, 2010). The available evidence suggests that the effects of ocean warming may occur throughout the year.  If winter temperatures exceed 12°C this is likely to inhibit germination, whilst summer temperatures greater than 20°C will cause mortality of Alaria esculenta.

Laminaria digitata is a polar to cold-temperate Atlantic species, which grows at temperatures of between 0-20 °C, with optimum growth at 10 °C (Bolton & Lüning, 1982), although both growth and photosynthesis show a significant decline at 18°C (Hargrave et al., 2017). Under experimental conditions, Laminaria digitata did not survive during 9 weeks of exposure to temperatures of 23°C (Wilson et al., 2015). During an exceptionally warm summer in Norway, Sundene (1964) reported mortality of sporophytes exposed to temperatures of 22-23°C.

Corallina officinalis may tolerate between -4 to 28°C (Lüning, 1990), although when Colthart & Johansen (1973) exposed this species to a number of different temperatures, they found that growth was maintained at 18°C and ceased at 25°C. Abrupt temperature changes (10°C in California, Seapy & Littler 1984; 4.8 to 8.5°C, Hawkins & Hartnoll, 1985) resulted in dramatic declines. However, in both cases recovery was rapid, suggesting that the crustose bases survived.

Sensitivity assessment. Alaria esculenta is competitively inferior to Laminaria digitata in the northeast Atlantic (Hawkins & Harkin, 1985) where it is confined to more exposed shores and can colonize sheltered environments only if Laminaria digitata is removed. As temperatures rise to up to 20°C, it is expected that the competitive ability of this species will decrease further, while as temperatures rise above 20°C both Alaria esculenta and Laminaria digitata will be lost, with more, warm adapted, algae taking their place. Brodie et al. (2014) predict that Alaria esculenta will be completely lost from Boreal habitats by the end of this century, whilst all kelp species will be lost from the Lusitanian region, replaced by smaller, fleshy algae.

Sea surface temperatures around the UK are currently between 6-19°C (Huthnance, 2010). Under the middle emission scenario, Alaria esculenta is likely to be lost from most of the UK, although it may manage to maintain some populations in Scotland, where current summer temperatures often reach 14°C, leading to potential summer temperatures of 17°C. Laminaria digitata is likely to be lost in southern parts of the UK, where summer temperatures will exceed 20°C, but will likely remain present around the rest of the UK. Corallina officinalis and other warm-tolerant species should be able to cope with temperature increases under a middle emission scenario. As Alaria esculenta is the main characterizing species for this study and is expected to be lost from a large portion of the UK, under this scenario resistance is assessed as ‘Low’. Resilience is assessed as ‘Very Low’, as the loss is likely to be a long-term decline, due to the long-term nature of ocean warming. Therefore, this biotope is assessed as having ‘High’ sensitivity to ocean warming under a middle emission scenario.

For the high emission scenario and extreme scenario, sea temperatures may rise by 4-5°C to give potential southern summer temperatures of 23-24°C and northern summer temperatures of 18-19°C. Under these scenarios, it is likely that Alaria esculenta will be lost almost completely from the UK, and Laminaria digitata will retreat northwards, with large losses across England, Ireland, and Wales. Therefore, resistance is assessed as ‘None’, and resilience is assessed as ‘Very low’. Overall, this biotope is assessed as having ‘High’ sensitivity to ocean warming for the high emission scenario and the extreme scenario.  

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 due to increased air-sea heat flux are predicted to occur more frequently, at an increased intensity and last for longer by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018). Marine heatwaves are known to cause significant impacts to kelp forests, particularly if a population is found towards the edge of its southern limit (Smale et al., 2019). In Baja California, Mexico, an extreme heat event between 2014 – 2016 led to both a decrease in the density of Macrocystis pyrifera and a decrease in the number of fronds per individual (Arafeh-Dalmau et al., 2019). There was also a significant change to the understory algal composition, and half the fish and invertebrates associated with this habitat disappeared. The same heatwave, coupled with a loss of starfish through disease and an increase in urchin grazing, led to the loss of > 90% of Macrocystis pyrifera from 350 km of coastline in northern California (Rogers-Bennett & Catton, 2019).

As this biotope occurs on the intertidal-subtidal margin, it will experience both elevated air temperatures and seawater temperatures during a heatwave, which may lead to doubly stressful conditions (Thomsen et al., 2019). Air temperatures tend to be more variable and extreme than seawater temperatures (Helmuth et al., 2002). While the south of the UK has a mean summer daily high temperature of 21°C, temperatures can often reach ≥30°C (Met_Office, 2016). Temperature loggers on the west coast of Scotland recorded intertidal temperatures on the high shore exceeding 40°C in 7 of the 11 years it was monitored (Burrows, 2017), showing the extreme temperatures that intertidal species have to cope with. Following the marine heatwave of 2017/2018 in New Zealand, the intertidal bull kelp, Durvillaea poha, was lost from part of the coastline (Thomsen et al., 2019). Air temperatures are likely to lead to enhanced desiccation (Thomsen et al., 2019), although this may be partially ameliorated by wave action in this biotope, as it only occurs on very exposed to moderately exposed shorelines.

Under experimental exposure to different seawater temperatures, germination in Alaria esculenta decreased at 16°C, whilst temperatures of 21°C were lethal to sporophytes. Similarly, when Laminaria digitata was exposed to typical spring-summer water temperatures (12, 16, 20°C), a predicted increase in summer temperature (23°C), and potential heatwave temperatures in shallow waters (26, 29°C) for nine weeks, no survival occurred at temperatures higher than 20°C (Wilson et al., 2015).

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 and 19°C in Scotland. Under the middle emission scenario, both Alaria esculenta and Laminaria digitata are likely to be lost from the southern parts of the UK (see Global warming pressure).  Any remaining populations of Alaria esculenta are likely to suffer severe mortality as a result of a heatwave of this magnitude. In Scotland, a significant proportion of Laminaria digitata is likely to survive in areas where temperatures do not exceed 20°C but will suffer mortality elsewhere. Therefore, resistance has been assessed as ‘None’. As a further heatwave is likely to affect this habitat before full recovery, recovery has been assessed as ‘Low.’ Therefore, this biotope is assessed as having ‘High’ sensitivity to marine heatwaves under the middle emission scenario.

Under the high emission scenario, if heatwaves occur every two years by the end of this century, reaching a maximum intensity of 3.5°C for 120 days, this could lead to the heatwave lasting the entire summer with temperatures reaching up to 26.5°C in southern England and 21.5°C in Scotland. Under the high emission scenario, Alaria esculenta is likely to be almost wiped out in the UK, whilst Laminaria digitata may be lost from large parts of the UK as its range contracts (see Global warming pressure). Therefore, resistance has been assessed as ‘None’. As widespread mortality may lead to a lack of viable sporophytes for recruitment, resilience has been assessed as ‘Very low.’ Therefore, this biotope is assessed as having ‘High’ sensitivity to marine heatwaves under the high emission scenario.

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 due to increased air-sea heat flux are predicted to occur more frequently, at an increased intensity and last for longer by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018). Marine heatwaves are known to cause significant impacts to kelp forests, particularly if a population is found towards the edge of its southern limit (Smale et al., 2019). In Baja California, Mexico, an extreme heat event between 2014 – 2016 led to both a decrease in the density of Macrocystis pyrifera and a decrease in the number of fronds per individual (Arafeh-Dalmau et al., 2019). There was also a significant change to the understory algal composition, and half the fish and invertebrates associated with this habitat disappeared. The same heatwave, coupled with a loss of starfish through disease and an increase in urchin grazing, led to the loss of > 90% of Macrocystis pyrifera from 350 km of coastline in northern California (Rogers-Bennett & Catton, 2019).

As this biotope occurs on the intertidal-subtidal margin, it will experience both elevated air temperatures and seawater temperatures during a heatwave, which may lead to doubly stressful conditions (Thomsen et al., 2019). Air temperatures tend to be more variable and extreme than seawater temperatures (Helmuth et al., 2002). While the south of the UK has a mean summer daily high temperature of 21°C, temperatures can often reach ≥30°C (Met_Office, 2016). Temperature loggers on the west coast of Scotland recorded intertidal temperatures on the high shore exceeding 40°C in 7 of the 11 years it was monitored (Burrows, 2017), showing the extreme temperatures that intertidal species have to cope with. Following the marine heatwave of 2017/2018 in New Zealand, the intertidal bull kelp, Durvillaea poha, was lost from part of the coastline (Thomsen et al., 2019). Air temperatures are likely to lead to enhanced desiccation (Thomsen et al., 2019), although this may be partially ameliorated by wave action in this biotope, as it only occurs on very exposed to moderately exposed shorelines.

Under experimental exposure to different seawater temperatures, germination in Alaria esculenta decreased at 16°C, whilst temperatures of 21°C were lethal to sporophytes. Similarly, when Laminaria digitata was exposed to typical spring-summer water temperatures (12, 16, 20°C), a predicted increase in summer temperature (23°C), and potential heatwave temperatures in shallow waters (26, 29°C) for nine weeks, no survival occurred at temperatures higher than 20°C (Wilson et al., 2015).

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 and 19°C in Scotland. Under the middle emission scenario, both Alaria esculenta and Laminaria digitata are likely to be lost from the southern parts of the UK (see Global warming pressure).  Any remaining populations of Alaria esculenta are likely to suffer severe mortality as a result of a heatwave of this magnitude. In Scotland, a significant proportion of Laminaria digitata is likely to survive in areas where temperatures do not exceed 20°C but will suffer mortality elsewhere. Therefore, resistance has been assessed as ‘None’. As a further heatwave is likely to affect this habitat before full recovery, recovery has been assessed as ‘Low.’ Therefore, this biotope is assessed as having ‘High’ sensitivity to marine heatwaves under the middle emission scenario.

Under the high emission scenario, if heatwaves occur every two years by the end of this century, reaching a maximum intensity of 3.5°C for 120 days, this could lead to the heatwave lasting the entire summer with temperatures reaching up to 26.5°C in southern England and 21.5°C in Scotland. Under the high emission scenario, Alaria esculenta is likely to be almost wiped out in the UK, whilst Laminaria digitata may be lost from large parts of the UK as its range contracts (see Global warming pressure). Therefore, resistance has been assessed as ‘None’. As widespread mortality may lead to a lack of viable sporophytes for recruitment, resilience has been assessed as ‘Very low.’ Therefore, this biotope is assessed as having ‘High’ sensitivity to marine heatwaves under the high emission scenario.

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), with it expected to drop up to a further 0.35 units by the end of this century, dependent on the emission scenario. Marine autotrophs will generally benefit from ocean acidification, through an increase in the availability of aqueous CO₂ for photosynthesis (Koch et al., 2013). Most species of kelp appear to be under-saturated in respect to carbon dioxide, although they can generally utilise HCO₃ and have external carbonic anhydrase for extracellular dehydration of HCO₃ to CO₂ (Koch et al., 2013).

Under experimental conditions, Iñiguez et al. (2016a) found that although photosynthesis remained stable in Alaria esculenta in response to increasing CO₂, the growth rate increased. Similarly, Gordillo et al. (2015) found heightened growth rates in Alaria esculenta when exposed to increased CO₂, although this increase was not significant and less pronounced than in Saccharina lattisima.

Research on other kelp species has revealed a positive or neutral effect of ocean acidification (Roleda et al., 2012, Fernández et al., 2015, Nunes et al., 2015, Iñiguez et al., 2016b, a), except for one study, which found that ocean acidification negatively impacted photosynthesis and growth in the southern hemisphere species, Ecklonia radiata (Britton et al., 2016). Whilst there is currently no research on the impact of ocean acidification on Laminaria digitata, it is likely that this species will either benefit or not be negatively impacted by ocean acidification.

Corallina officinalis is a highly calcified, erect, red algae. Results of experimental CO₂ enrichment suggest that this species could be significantly negatively affected by future ocean acidification. Hofmann et al. (2012) found that growth and photosynthesis decreased as a result of a 0.3 unit decrease in pH. Further investigation showed that skeletal CaCO₃ decreased with increasing CO₂ at levels expected for both the middle emission and high emission scenarios, although this decrease was small (< 2%) (Hofmann et al., 2013). Yildiz et al. (2013) showed that although CaCO₃ decreased in Corallina officinalis as a result of ocean acidification, photosynthesis increased. When ocean acidification was combined with an increase in UV radiation, this led to an increase in growth rate. They summarised that a decrease in CaCO₃ content may not be negative but may lead to this species absorbing and using light differently.

Sensitivity assessment. Kelp forests occur in a naturally variable pH habitat, with diel fluctuations of 0.3 - 0.45 pH units (Krause-Jensen et al., 2015, Britton et al., 2016), and boundary layer pH fluctuation of up to 0.8 units (Krause-Jensen et al., 2015). The kelps Alaria esculenta and Laminaria digitata are not expected to suffer negative impacts from future acidification, and whilst calcium carbonate content of Corallina officinalis may reduce as a result of acidification, this species is expected to survive future CO₂ enrichment of the oceans. Therefore, under both the middle and high emission scenario resistance is assessed as ‘High’, and resilience is assessed as ‘High’ leading to a score of ‘Not sensitive’.

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), with it expected to drop up to a further 0.35 units by the end of this century, dependent on the emission scenario. Marine autotrophs will generally benefit from ocean acidification, through an increase in the availability of aqueous CO₂ for photosynthesis (Koch et al., 2013). Most species of kelp appear to be under-saturated in respect to carbon dioxide, although they can generally utilise HCO₃ and have external carbonic anhydrase for extracellular dehydration of HCO₃ to CO₂ (Koch et al., 2013).

Under experimental conditions, Iñiguez et al. (2016a) found that although photosynthesis remained stable in Alaria esculenta in response to increasing CO₂, the growth rate increased. Similarly, Gordillo et al. (2015) found heightened growth rates in Alaria esculenta when exposed to increased CO₂, although this increase was not significant and less pronounced than in Saccharina lattisima.

Research on other kelp species has revealed a positive or neutral effect of ocean acidification (Roleda et al., 2012, Fernández et al., 2015, Nunes et al., 2015, Iñiguez et al., 2016b, a), except for one study, which found that ocean acidification negatively impacted photosynthesis and growth in the southern hemisphere species, Ecklonia radiata (Britton et al., 2016). Whilst there is currently no research on the impact of ocean acidification on Laminaria digitata, it is likely that this species will either benefit or not be negatively impacted by ocean acidification.

Corallina officinalis is a highly calcified, erect, red algae. Results of experimental CO₂ enrichment suggest that this species could be significantly negatively affected by future ocean acidification. Hofmann et al. (2012) found that growth and photosynthesis decreased as a result of a 0.3 unit decrease in pH. Further investigation showed that skeletal CaCO₃ decreased with increasing CO₂ at levels expected for both the middle emission and high emission scenarios, although this decrease was small (< 2%) (Hofmann et al., 2013). Yildiz et al. (2013) showed that although CaCO₃ decreased in Corallina officinalis as a result of ocean acidification, photosynthesis increased. When ocean acidification was combined with an increase in UV radiation, this led to an increase in growth rate. They summarised that a decrease in CaCO₃ content may not be negative but may lead to this species absorbing and using light differently.

Sensitivity assessment. Kelp forests occur in a naturally variable pH habitat, with diel fluctuations of 0.3 - 0.45 pH units (Krause-Jensen et al., 2015, Britton et al., 2016), and boundary layer pH fluctuation of up to 0.8 units (Krause-Jensen et al., 2015). The kelps Alaria esculenta and Laminaria digitata are not expected to suffer negative impacts from future acidification, and whilst calcium carbonate content of Corallina officinalis may reduce as a result of acidification, this species is expected to survive future CO₂ enrichment of the oceans. Therefore, under both the middle and high emission scenario resistance is assessed as ‘High’, and resilience is assessed as ‘High’ leading to a score of ‘Not sensitive’.

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). Sea-level rise is expected to lead to substantial loss of intertidal habitats. Rocky shores backed by cliffs constitute about 80% of oceanic coastlines globally and in Britain, 42% of the coastline is hard rock, with many areas having cliffs behind the shore (Jackson & McIlvenny, 2011).

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

Sensitivity assessment. This biotope is found on steep and vertical rock on the lower shore and shallow subtidal, and therefore an increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope. Beds may be able to expand their range and migrate upwards to compensate for sea-level rise, if not constrained by lack of suitable habitat (IPCC, 2019). In this assessment we have assessed on a worst-case-scenario basis, assuming that landward migration is not possible, which is likely to lead to the depth distribution of Alaria esculenta shrinking significantly in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery. Alaria esculenta is generally limited to a depth of 1-2 m, as below this depth it is usually outcompeted by Laminaria hyperborea. As depth increases due to sea-level rise, Laminaria hyperborea is expected to extend its distribution shorewards.

Under the middle emission, high emission and extreme scenarios (50, 70 and 107 cm sea-level rise), it is expected that between 25 - 75% of this biotope is likely to be lost.  Therefore, for all three scenarios, resistance has been assessed as ‘Low’, and resilience as ‘Very low’.  Therefore, sensitivity is assessed as ‘High’.

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). Sea-level rise is expected to lead to substantial loss of intertidal habitats. Rocky shores backed by cliffs constitute about 80% of oceanic coastlines globally and in Britain, 42% of the coastline is hard rock, with many areas having cliffs behind the shore (Jackson & McIlvenny, 2011).

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

Sensitivity assessment. This biotope is found on steep and vertical rock on the lower shore and shallow subtidal, and therefore an increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope. Beds may be able to expand their range and migrate upwards to compensate for sea-level rise, if not constrained by lack of suitable habitat (IPCC, 2019). In this assessment we have assessed on a worst-case-scenario basis, assuming that landward migration is not possible, which is likely to lead to the depth distribution of Alaria esculenta shrinking significantly in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery. Alaria esculenta is generally limited to a depth of 1-2 m, as below this depth it is usually outcompeted by Laminaria hyperborea. As depth increases due to sea-level rise, Laminaria hyperborea is expected to extend its distribution shorewards.

Under the middle emission, high emission and extreme scenarios (50, 70 and 107 cm sea-level rise), it is expected that between 25 - 75% of this biotope is likely to be lost.  Therefore, for all three scenarios, resistance has been assessed as ‘Low’, and resilience as ‘Very low’.  Therefore, sensitivity is assessed as ‘High’.

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). Sea-level rise is expected to lead to substantial loss of intertidal habitats. Rocky shores backed by cliffs constitute about 80% of oceanic coastlines globally and in Britain, 42% of the coastline is hard rock, with many areas having cliffs behind the shore (Jackson & McIlvenny, 2011).

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

Sensitivity assessment. This biotope is found on steep and vertical rock on the lower shore and shallow subtidal, and therefore an increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope. Beds may be able to expand their range and migrate upwards to compensate for sea-level rise, if not constrained by lack of suitable habitat (IPCC, 2019). In this assessment we have assessed on a worst-case-scenario basis, assuming that landward migration is not possible, which is likely to lead to the depth distribution of Alaria esculenta shrinking significantly in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery. Alaria esculenta is generally limited to a depth of 1-2 m, as below this depth it is usually outcompeted by Laminaria hyperborea. As depth increases due to sea-level rise, Laminaria hyperborea is expected to extend its distribution shorewards.

Under the middle emission, high emission and extreme scenarios (50, 70 and 107 cm sea-level rise), it is expected that between 25 - 75% of this biotope is likely to be lost.  Therefore, for all three scenarios, resistance has been assessed as ‘Low’, and resilience as ‘Very low’.  Therefore, sensitivity is assessed as ‘High’.

Temperature increase (local)

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

Evidence

Alaria esculenta is a northern/boreal species that has been recorded from Brittany, France to Northern Norway (Birkett et al., 1998b). Sea temperature regulates metabolism and reproduction and defines the regional distribution of Alaria esculenta (Fredersdorf et al., 2009).  The southern limit of Alaria esculenta has been defined at the 20°C isotherm (Munda & Lüning, 1977; Fredersdorf et al., 2009), however, it is common north of the 16°C isotherm (Munda & Lüning, 1977). As a result of this upper temperature threshold, Alaria esculenta is largely absent from the southern North Sea and English channel where summer temperatures can exceed 16°C.

Munda & Lüning (1977) observed temperatures of 16-17°C sustained over 2 weeks in Helgoland, Germany, were lethal to resident Alaria esculenta. Experimental observations showed that acute exposure to ≥21°C is lethal to Alaria esculenta causing bleaching and disintegration (Sundene, 1962; Fredersdorf et al., 2009). At its northern range edge (Svalbard) it is a prominent macroalga on sublittoral fringe bedrock.  At these latitudes average, summer temperature can reach 5°C, with an average annual sea temperature of 3°C (1980-2014, Beszczynska-Möller & Dye, 2013). Experimental observations conducted by Fredersdorf et al. (2009) found the optimal temperature for sporophyte photosynthesis was within the range of 13-17°C, however, the optimal temperature for Alaria esculenta germination is 2-12°C (Fredersdorf et al., 2009).

Alaria esculenta has an approximate mid-range within southern Norway (60 deg 65 deg North) (Birkett et al., 1998b), and as such IR.HIR.KFaR.Ala and IR.HIR.KFaR.AlaAnCrSp (plus associated sub-biotopes) have a southerly distribution when considering the geographic distribution of Alaria esculenta. Throughout the UK northern to southern Sea Surface Temperature (SST) ranges from 8-16°C in summer and 6-13°C in winter (Beszczynska-Möller & Dye, 2013). The available evidence suggests that the effects of an increase in temperature would be seasonally variable, with higher impacts during periods of spore release (Nov-march) and germination. A 5°C increase in temperature for one month may cause high mortality, limit photosynthetic ability plus germination rates.  A 2°C increase in temperature for one year may limit germination; however sporophyte photosynthetic ability may not be dramatically affected. Temperature increases of 2-5°C at the southern extreme of Alaria esculenta’ range (Brittany, France) is likely to cause high mortality.

Corallina officinalis may tolerate between minus 4 to 28°C (Lüning, 1990). Abrupt temperature changes (10°C in California, Seapy & Littler 1984; 4.8 to 8.5°C, Hawkins & Hartnoll, 1985) resulted in dramatic declines. However, in both cases recovery was rapid, suggesting that the crustose bases survived. Therefore, both Alaria esculenta and Corallina officinalis are probably intolerant of acute short-term temperature change of 5°C for a month. Long-term change of 2°C may reduce the southern limit of the population of Alaria esculenta.

Sensitivity assessment. Resistance to the pressure is considered ’None‘, and resilience ’High‘. The sensitivity of this biotope to an increase in temperature has been assessed as ’Medium‘. This sensitivity assessment takes into account a one month temperature increase of 5°C. The effects of a 2°C increase in temperature for one year is likely to have less of an impact. In the later scenario, resistance would be assessed as 'Medium', and resilience 'High'. Sensitivity would be 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

Alaria esculenta is a northern/boreal species that has been recorded from Brittany, France to Northern Norway (Birkett et al., 1998b). Sea temperature has been cited as an influential abiotic stressor; responsible for regulating metabolism and reproduction, plus defining the regional distribution of Alaria esculenta (Fredersdorf et al., 2009). At Alaria’snorthern range edge (Svalbard) it is a prominent macro-algae on sublittoral fringe bedrock. At these latitudes, average summer temperature can reach 5°C, and average annual sea temperature 3°C (1980-2014, Beszczynska-Möller & Dye, 2013). Experimental observations conducted by Fredersdorf et al., (2009) found the optimal temperature for sporophyte photosynthesis was within the range of 13-17°C, however, the optimal temperatures for Alaria esculenta germination is 2-12°C (Fredersdorf et al., 2009).

Alaria esculenta has an approximate mid-range within southern Norway (60 deg to 65 deg North) (Birkett et al., 1998b), and as such IR.HIR.KFaR.Ala and IR.HIR.KFaR.AlaAnCrSp (plus associated sub-biotopes) have a southerly distribution when considering the geographic distribution of Alaria esculenta. Throughout the UK northern to southern Sea Surface Temperature (SST) ranges from 8-16°C in summer and 6-13°C in winter (Beszczynska-Möller & Dye, 2013). A 5°C decrease in temperature for one month at Alaria esculenta’ approximate mid-range may affect the photosynthetic ability of sporophytes, however not impact germination and hence recruitment. A 2°C increase in temperature for one year at Alaria esculenta’ approximate mid-range is not likely to significantly affect Alaria esculenta.

Sensitivity assessment. Resistance to the pressure is considered ’High‘, and resilience ’High‘. The sensitivity of this biotope to an increase in temperature has been assessed as ’Not Sensitive‘. 

Salinity increase (local)

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

Evidence

Lüning (1990) suggest that kelps are stenohaline, their general tolerance to salinity as a phenotypic group covering 16-50 psu over a 24 hr period. Optimal growth probably occurs between 30-35 psu (MNCR category-Full Salinity) and growth rates are likely to be affected by periodic salinity stress.

Karsten (2007) tested the photosynthetic ability of Alaria esculenta under acute 2 and 5 day exposure to salinity treatments ranging from 5-60 psu. A control experiment was also carried at 34 PSU. Between 10-50 psu Alaria esculenta showed high photosynthetic ability at 83-94% of the control. Hypersaline treatments with 55-60 psu led to a 30% reduction in photosynthetic ability, ~70% of the control level. At 5 psu Alaria esculenta showed a low photosynthetic ability at 15.8% of the control. After 5 days at 5 psu, all Alaria esculenta specimens were bleached and none survived. Karsten (2007) suggested that Alaria esculenta photosynthetic ability is highly affected by acute exposure to hyposaline conditions (<10 psu). The effect of long-term salinity changes (>5 days) or the effect of salinity >60 psu on Alaria esculenta’ photosynthetic ability was not tested. The experiment was conducted in the Arctic, and the authors suggest that at extremely low water temperatures (1-5°C) macro-algal acclimation to rapid salinity changes could be slower than at temperate latitudes. It is, therefore, possible that Alaria esculenta maybe be able to acclimate to salinity changes more effectively and quicker in UK waters, however, evidence for this is limited.

Corallina officinalis is restricted to full salinity waters in the Baltic and grows maximally between 33 and 38 psu in Texan lagoons (Kinne 1971a). This biotope is likely to be exposed to short-term freshwater runoff at low tide but is likely to be intolerant of long-term changes in salinity, which are likely to depress its upper limit and reduce the extent of the population.

Sensitivity assessment. IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes have been recorded exclusively in full salinity (30-40‰) (Connor et al., 2004). Karsten (2007) suggests that at salinities ranging from 10-50 psu Alaria esculenta photosynthetic ability was high. At salinities >50 psu, photosynthetic ability was reduced by 30% but no mortality of the specimens was recorded. Resistance to the pressure is considered ’Medium‘, as other characterizing species (e.g. sponges, ascidians) are likely to be more sensitive to hypersaline conditions and resilience ’High’. The sensitivity of this biotope to an increase in salinity 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

Lüning (1990) suggest that kelps are stenohaline, their general tolerance to salinity as a phenotypic group covering 16-50 psu over a 24 hr period. Optimal growth probably occurs between 30-35 psu (MNCR category-Full Salinity) and growth rates are likely to be affected by periodic salinity stress.

Karsten (2007) tested the photosynthetic ability of Alaria esculenta under acute 2 and 5 day exposure to salinity treatments ranging from 5-60 psu. A control experiment was also carried at 34 psu. Between 10-50 psu Alaria esculenta showed high photosynthetic ability at 83-94% of the control. Hypersaline treatments with 55-60 psu led to a 30% reduction in photosynthetic ability, about 70% of the control level. At 5 psu Alaria esculenta showed a low photosynthetic ability at 15.8% of the control. After 5 days at 5 psu, all Alaria esculenta specimens were bleached and none survived. Karsten (2007) suggested that Alaria esculenta photosynthetic ability is highly affected by acute exposure to hyposaline conditions (<10 psu). The effect of long-term salinity changes (>5 days) or the effect of salinity >60 psu on Alaria esculenta’ photosynthetic ability was not tested. The experiment was conducted in the Arctic, and the authors suggest that at extremely low water temperatures (1-5°C) macro-algal acclimation to rapid salinity changes could be slower than at temperate latitudes. It is, therefore, possible that Alaria esculenta maybe be able to acclimate to salinity changes more effectively and quicker in UK waters, however, evidence for this is limited.

Corallina officinalis is restricted to full salinity waters in the Baltic and grows maximally between 33 and 38 psu in Texan lagoons (Kinne 1971a). This biotope is likely to be exposed to short-term freshwater runoff at low tide but is likely to be intolerant of long-term changes in salinity, which are likely to depress its upper limit and reduce the extent of the population.

Sensitivity assessment. IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes have been recorded exclusively in full salinity (30-40‰) (Connor et al., 2004). Karsten (2007) suggests that at salinities ranging from 10-50 PSU Alaria esculenta photosynthetic ability was high. At 5 PSU Alaria esculenta showed a dramatic decline in photosynthetic ability and after 5 days specimens bleached and did not survive. Sundene (1962) also noted that Alaria esculenta sporophytes grew poorly below 25 PSU. A decrease of one MNCR salinity scale to “Reduced Salinity” (18-30‰) may reduce growth rates, however not cause high mortality of Alaria esculenta. Resistance to the pressure is therefore considered ’Medium‘, as other characterizing species (e.g. sponges, ascidians) are likely to be more sensitive to hyposaline conditions, and resilience ’High‘.  The sensitivity of this biotope to an increase in salinity 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

Alaria esculenta dominates the sublittoral fringe in areas exposed to severe wave action or where water surges along the sides of gullies/steep bedrock faces (Lewis, 1964; Connor et al., 2004). The high wave exposure that defines IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes damages other laminarians and generally excludes them.  In less wave exposed locations Alaria esculenta is out-competed by other Laminarians, e.g. Laminaria digitata and Laminaria hyperborea (Connor et al., 2004). IR.HIR.KFaR.AlaAnCrSp and IR.HIR.KFaR.Ala plus associated sub-biotopes are recorded within moderately strong (0.5-1.5 m/sec)-weak (<0.5 m/sec) tidal streams, but have been recorded in very strong (>3 m/sec) tidal streams. Therefore, elevated tidal flows (>3 m/sec) may increase Alaria esculenta dislodgment (Birkett et al., 1998b).

Increased tidal flow may remove fronds of Corallina officinalis however calcification is thought to be an adaptation to mechanical damage (Littler & Kauker, 1984). Increases in water flow rate may facilitate the colonization of filter feeding organisms within the understorey and IR.HIR.KFaR.Ala.Myt may dominate over IR.HIR.KFaR.Ala.Ldig. Decreases in water flow are likely to have the opposite effect (Connor et al. 2004). Changes in the water flow regimes under kelp canopies can modify larval supply and settlement (Eckman, 1983), and affect the growth and survival of Mytillus edulis (Eckman & Duggins, 1991). Mytillus edulis settlement has been found significantly higher in close proximity to Alaria esculenta and is thought to increase beneath the canopy (Bégin et al., 2004). Therefore any loss of Alaria esculenta, as a result of changes to local water movements, may affect Mytilus edulis recruitment.

Sensitivity assessment. IR.HIR.KFaR.AlaAnCrSp and IR.HIR.KFaR.Ala plus associated sub-biotopes are found in a wide range of tidal flows but exclusively in wave disturbed areas, which generally exclude other laminarians. Changes in tidal flow are not likely to independently affect the dominance of Alaria esculenta, however, may affect the understorey community.  Nevertheless, wave exposure is the dominant source of water movement in theses biotope, and a change in water flow of 0.1-0.2 m/s is unlikely to be significant. Therefore, resistance has been assessed as ’High‘ and resilience ’High‘. 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

An increase in emergence will result in an increased risk of desiccation. Increased immersion may allow IR.HIR.KFaR.Ala biotopes to extend higher up the shore. However, Alaria esculenta forest will come under increased competition from Laminaria hyperborea in the shallow infralittoral. In this scenario IR.HIR.KFaR.Ala biotope distribution may shift on the shore, however, biotope structure will remain.

Alaria esculenta may extend into the lower eulittoral in extremely wave exposed conditions. However, these marginal populations have a reduced age range in comparison to subtidal populations due to desiccation increasing mortality of Alaria esculenta at low tide. An increase in desiccation is likely to remove Alaria esculenta. The resultant loss of canopy would expose Corallina officinalis turf and macrofaunal crust to desiccation and/or damage by high light intensity (bleaching). Hawkins & Harkin (1985) noted that encrusting corallines and Corallina officinalis often die when their protective algal canopy is removed. Severe damage was noted in Corallina officinalis as a result of unusually hot and sunny weather in the UK summer 1983 (Hawkins & Hartnoll, 1985). Laminaria digitata is likely to be intolerant of desiccation and destruction of its meristem (base of the blade), caused by increased wave action at low tide, will kill the sporophyte. Therefore, both IR.HIR.KFaR.Ala.Myt and IR.HIR.KFaR.Ala.Ldig are likely to be highly intolerant of increases in desiccation and the upper limit of the population would be depressed. Desiccation is unlikely to be relevant in IR.HIR.KFaR.AlaAnCrSp due to its depth (15-35 m BCD) (Connor et al., 2004).

Sensitivity assessment. Resistance to this pressure is considered ’Low‘, and resilience ’High‘. The sensitivity of this biotope to a change in emergence is considered as ’Low‘.  

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

Alaria esculenta dominates the sublittoral fringe in areas exposed to severe wave action or where water surges along the sides of gullies (Lewis, 1964). A decrease in local wave height will increase spatial competition from other laminarians (Connor et al., 2004). Increased wave exposure may remove fronds of Corallina officinalis however calcification is thought to an adaptation to mechanical damage (Little & Kauker, 1984) and the fronds grow as a compact (short) turf in wave exposed conditions.

IR.HIR.KFaR.AlaAnCrSp occurs at one site, Rockall, Scotland where extreme oceanic swell excludes Laminaria hyperborea in the infralittoral from 14-35 m. IR.HIR.KFaR.Ala.Myt occurs predominantly on sublittoral fringe bedrock in very exposed to exposed wave exposure.  Extremely wave exposed variants of IR.HIR.KFaR.Ala.Myt can extend to 15 m BCD where Alaria esculenta replaces Laminaria hyperborea as the assemblage dominant, and Mytillus edulis is a common understorey species in the sublittoral fringe variant (Bégin et al., 2004, Connor et al., 2004) but as depth increases Tubularia spp,. becomes more abundant. IR.HIR.KFaR.Ala.Ldig occurs predominately at exposed-moderately wave exposed sites, where Laminaria digitata can spatially compete with Alaria esculenta (Connor et al., 2004).

Sensitivity assessment. The abundance of Alaria esculenta is highly affected by the degree of wave exposure at a site. Within IR.HIR.KFaR.Ala, increasing wave exposure may favour IR.HIR.KFaR.Ala.Myt over IR.HIR.KFaR.Ala.Ldig (Connor et al. 2004). Further increases in wave exposure may cause damage to Laminaria hyperborea, allowing Alaria esculenta to dominate the infralittoral. Kelp clearance experiments have shown that at moderate or lower wave exposure sites Laminaria digitata can out-compete Alaria esculenta so that a decrease in wave exposure is likely to result in loss of the Alaria dominated biotopes. Alaria dominated biotopes are, therefore, sensitive to any activity or event that reduces incident wave energy.  However, a change of 3-5% in significant wave height (the benchmark) is unlikely to be significant in the wave exposed conditions favoured by theses biotopes.  Therefore, resistance is recorded as ‘High’, with a ‘High’ resilience, resulting is an assessment of ‘Not sensitive’ at the benchmark level.

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.

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including metal contamination) affected six marine habitats (including intertidal and subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types. Johnston & Roberts (2009) also highlighted that macroalgal communities are relatively tolerant to contamination but that metal and nutrient impacted intertidal communities can have low diversity assemblages which are dominated by opportunistic and fast growing species (Johnston & Roberts, 2009).

Mercury (organic > inorganic) is highly toxic to macrophytes (Bryan, 1984; Cole et al., 1999). Mercury and copper were lethal at 0.05 mg/l and 0.1 mg/l respectively and toxic at 0.05 mg/l and 0.01 mg/l respectively in Laminaria hyperborea. Zinc and Cadmium were lethal at 5 mg/l and 10 mg/l respectively. The presence of alginates in kelp tissue is thought to sequester heavy metals in a biologically unavailable form. It is likely that laminarians such as Alaria esculenta are relatively tolerant of heavy metals except at high concentrations at high levels. Little information on heavy metal tolerance of corallines was found.

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.

The mucilaginous coating on kelp fronds is thought to protect them from coatings of oil. Hydrocarbons in solution reduce photosynthesis and may be algicidal. Reduction in photosynthesis is dependent on the type of oil, its concentration and length of exposure, oil-water mixture and irradiance in experimental trials (Lobban & Harrison, 1994). Subtidal populations are only exposed to oil emulsions or oil adsorbed particles. Kelps are relatively insensitive to dispersants (Birkett et al., 1998b) e.g. Laminaria digitata exposed to diesel oil at 0.130 mg/l reduced growth by 50% in a 2 year experiment. No growth inhibition was noted at 0.03 mg/l and the plants recovered completely in oil free conditions. Corallina officinalis, however, exhibited dramatic bleaching after the Sea Empress oil spill and died after the Torrey Canyon spill (Crump et al., 1999; Smith, 1968). Encrusting corallines and Corallina officinalis recovered from the Sea Empress spill quickly, bleaching only affecting the fronds or surface of crustose forms. Grazing gastropods, e.g. limpets are highly intolerant of oil spillage and if not killed are narcotized and washed offshore and/or consumed by predators. The lower littoral populations are likely to be most vulnerable to an oil spill and sublittoral fringe would be particularly affected at low tide. Although Alara esculenta may not be affected severely, the articulated coralline turf may be lost but recover quickly although the red algae may be intolerant. Grazers such as limpets, barnacles and meiofaunal crustaceans may also be lost from the community.

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.

Cole et al. (1999) suggested that macrophytes are generally intolerant of herbicides such as atrazine, simazine, diuron and linuron. For example, atrazine was lethal to Laminaria hyperborea sporophytes at 1mg/l and suppressed growth at 0.01 mg/l (Hopkin & Kain, 1978). Smith (1968) noted that Corallina officinalis was killed in areas of heavy spraying after the Torrey Canyon oil spill and affected at 6 m depth in areas of high wave action. High water specimens were more affected than low water specimens, presumably because they are emmersed for longer and had more contact with oil and dispersants. Gastropods are known to be highly sensitive to endocrine disrupters such as TBT. Crustaceans (e.g. amphipods, isopods, ostracods, copepods and barnacles) are also susceptible to endocrine disruption by synthetic chemicals. It is, therefore, likely that some taxa within IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes, especially grazing invertebrates and meiofauna will be intolerant of synthetic chemical contamination.

Radionuclide contamination

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

Evidence

No Evidence

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

Reduced oxygen concentrations have been shown to inhibit both photosynthesis and respiration in macroalgae (Kinne, 1977). Despite this, macroalgae are thought to buffer the environmental conditions of low oxygen, thereby acting as a refuge for organisms in oxygen depleted regions especially if the oxygen depletion is short-term (Frieder et al., 2012).  A rapid recovery from a state of low oxygen is expected if the conditions are transient, which is likely given the wave exposed distribution of defines IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes. If levels do drop below 4 mg/l negative effects on these organisms can be expected with adverse effects occurring below 2mg/l (Cole et al., 1999). Reduced oxygen levels are likely to inhibit photosynthesis and respiration but not cause a loss of the macroalgae population directly.  However, small invertebrate epifauna may be lost, causing a reduction in species richness.

Sensitivity assessment.  Due to the mixing experienced in strongly wave exposed environments, resistance has been assessed as “High” resilience as “High”. Sensitivity has been assessed as “Not Sensitive” at the pressure benchmark level.

Nutrient enrichment

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

Evidence

Organic enrichment is associated with eutrophication, increased siltation and turbidity (Fletcher, 1996). Eutrophication is associated with loss of perennial algae and replacement by mussels or opportunistic algae (Fletcher, 1996). Johnston & Roberts (2009) conducted a meta-analysis that reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including intertidal and subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types. Johnston & Roberts (2009) also highlighted that macroalgal communities are relatively tolerant to contamination, but that metal and nutrient impacted intertidal communities can have low diversity assemblages which are dominated by opportunistic and fast growing species (Johnston & Roberts, 2009). However due to the high wave exposure that defines IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes, it is likely that additional organic input to the system may be dispersed out of the biotope’s local vicinity (Johnston & Roberts, 2009). Increased nutrients may favour Mytilus edulis in IR.HIR.KFaR.Ala.Myt which may increase in cover and abundance. Corallina officinalis is also tolerant of polluted waters (Kindig & Littler, 1980).

Sensitivity assessment. IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes are however considered “Not Sensitive” at the benchmark level, which assumes compliance with good status as defined by the WFD.

Organic enrichment

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

Evidence

Organic enrichment is associated with eutrophication, increased siltation and turbidity (Fletcher, 1996). Eutrophication is associated with loss of perennial algae and replacement by mussels or opportunistic algae (Fletcher, 1996). Johnston & Roberts (2009) conducted a meta-analysis that reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected six marine habitats (including intertidal and subtidal reefs).  A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types. Johnston & Roberts (2009) also highlighted that macroalgal communities are relatively tolerant to contamination, but that contaminated intertidal communities can have low diversity assemblages which are dominated by opportunistic and fast growing species (Johnston & Roberts, 2009). Due to the high wave exposure that defines IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes, it is likely that additional organic input to the system may be dispersed out of the biotope’s local vicinity (Johnston & Roberts, 2009). Increased nutrients may favour Mytilus edulis in IR.HIR.KFaR.Ala.Myt which may increase in cover and abundance. Corallina officinalis is also tolerant of polluted waters (Kindig & Littler, 1980).

Sensitivity assessment. Resistance has been assessed as “Medium”, (to represent potential changes in species diversity), resilience as “High”. 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 substrata were replaced with sedimentary substrata this would represent a fundamental change in habitat type, which Alaria esculenta would not tolerate (Birkett et al., 1998b). The biotope would be lost.

Sensitivity assessment. Resistance to the pressure is considered ’None‘, and resilience ’Very low‘ or ‘None’. The sensitivity of this biotope to change from hard rock or artificial substrata to sedimentary or soft rock substrata  is 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 to hard rock biotopes.

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

Not relevant to hard rock biotopes.

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

The sublittoral fringe is unlikely to be significantly impacted by trampling due to its position of the lower shore but may be prone to abrasion from moorings or low tide landings. Given its resilience to wave action Alaria esculenta is unlikely to be significantly damaged by abrasion although the understorey coralline turf may suffer some damage. The coralline turf meiofauna will probably be lost as a result of trampling. Moderate trampling on articulated coralline algal turf in the New Zealand intertidal (Brown & Taylor, 1999; Schiel & Taylor, 1999) resulted in reduced turf height, declines in turf densities, and loss of crustose bases in some case probably due to loss of the canopy algae and resultant desiccation. Calcification is thought to an adaptation to grazing and sediment scour (Littler & Kauker, 1984).

If exposed to moorings, groundings, or passing fishing gear, the resultant abrasion may result in the physical removal of a proportion of the Alaria esculenta canopy. Depending on the scale of the impact, although no evidence of this impact was found.  However, Alaria esculenta has been shown to be an opportunistic colonizing species, capable of rapid recovery (see resilience section).

Sensitivity assessment. Resistance has been assessed as ’Medium‘, resilience as  ‘High‘. Sensitivity has been assessed as ’Low’.

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

Not 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

Suspended Particle Matter (SPM) concentration has a linear relationship with sub surface light attenuation (Kd) (Devlin et al., 2008). Light penetration influences the maximum depth at which kelp species can grow and it has been reported that laminarians grow at depths at which the light levels are reduced to 1 percent of incident light at the surface. Maximal depth distribution of laminarians therefore varies from 100 m in the Mediterranean to only 6-7 m in the silt laden German Bight. In European Atlantic waters the depth limit is typically 35 m.

Alaria esculenta is not found in areas of siltation and sediment scour (Birkett et al. 1998b). Increased siltation and sediment scour inhibits photosynthesis and algal growth, interfere with spore or larval recruitment plus smother germlings and gametophytes (Fletcher, 1996). However, the high degree of wave exposure that typically defines IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes is likely to clear suspended sediments relatively quickly. If low water clarity is persistent and wave exposure decreased then low energy silted kelp biotopes (IR.LIR.K) may proliferate. Once siltation returns to its pre-effect level the biotope is likely to recover its canopy within a year and the rest of the community in no more than five years. Increased siltation will also increase turbidity. Increased sediment my benefit Mytilus edulis and its abundance may increase in IR.HIR.KFaR.Ala.Myt although large individuals are likely to be removed by wave action.

Increased turbidity is likely to reduce the depth to which Alaria  esculenta can grow. However, an increase of one level in WFD water clarity scale for a period of one year is unlikely to affect the population since Alaria esculenta’s lower limit, is generally determined by competition from other Laminarians rather than light penetration.

Sensitivity assessment. Resistance has been assessed as Medium, Resilience as High. Sensitivity has been assessed as Low.

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

Due to their size juvenile sporophytes, germlings, gametophytes and spores are likely to be inundated by deposition of 5cm during a discrete event but the high wave exposure that defines the distribution of IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala (plus associated sub-biotopes deposited sediments) are likely to be removed rapidly and any effects of inundation are likely to be temporary.

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

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

Due to their size juvenile sporophytes, germlings, gametophytes and spores are likely to be inundated by deposition of 30cm during a discrete event but the high wave exposure that defines the distribution of IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala (plus associated sub-biotopes deposited sediments) are likely to be removed rapidly and any effects of inundation are likely to be temporary.

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

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

No evidence

Underwater noise changes

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

Evidence

Not relevant

Introduction of light or shading

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

Evidence

There was no evidence to suggest that anthropogenic light sources would affect IR.HIR.KFaR.AlaAnCrSp & IR.HIR.KFaR.Ala plus associated sub-biotopes. Shading (e.g. by construction of a pontoon, pier etc) could adversely affect the biotope in areas where the water clarity is also low, and tip the balance to shade tolerant species, resulting in the loss of the biotope directly within the shaded area, or a reduction in laminarian abundance from forest to park type biotopes.

Sensitivity assessment. Resistance is probably 'Low', with a 'High' resilience and a sensitivity of 'High', albeit with 'low' confidence due to the lack of direct evidence.

Barrier to species movement

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

Evidence

Not relevant. This pressure is considered applicable to mobile species, e.g. fish and marine mammals rather than seabed habitats. Physical and hydrographic barriers may limit the dispersal of spores but spore dispersal is not considered under the pressure definition and benchmark.

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.  Collision from grounding vessels is addressed under abrasion above.

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

No evidence was found

Introduction or spread of invasive non-indigenous species

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

Evidence

Competition with invasive macroalgae may be a potential threat to this biotope. Potential invasives include Undaria pinnatifida and Sargassum muticum. Sargassum muticum is a circumglobal invasive species and is recorded from Norway to Morocco and into the Mediterranean in the eastern Atlantic, from Alaska to Baja California in the eastern Pacific and from southern Russia to southern China in the western Pacific (Engelen et al., 2015). It colonizes a variety of habitats and can tolerate -1°C to 30°C and survive salinities below 10 ppt. its distribution is limited by the availability of hard substratum (e.g., stones >10 cm) and light (Staeher et al., 2000; Strong & Dring 2011; Engelen et al., 2015). It is most abundant between 1 and 3 m below mean water. It is a poor competitor under low light and only develops dense canopies in shallow areas (Engelen et al., 2015). In Limfjorden, Denmark between 1984 and 1997 (Staehr et al., 2000; Engelen et al., 2015; de Bettignies et al., 2021). In Limfjorden, the distribution of Sargassum muticum was limited to areas with hard substratum, in particular stones >10 cm in diameter, while smaller stones, gravel and sand were unsuitable. It was most abundant between 1 and 4 m in depth but had low cover at 0-0.5 m or 4-6 m, in the turbid waters of the Limfjorden. Limfjorden is wave sheltered but wave exposure has been reported to restrict the growth and survival of Sargassum muticum (Staehr et al., 2000). 

Undaria pinnatifida (Wakame or Asian kelp) is a large brown seaweed and an Invasive Non-Indigenous Species (INIS) that could out-compete native UK kelp species (see Farrell & Fletcher, 2006; Thompson & Schiel, 2012; Brodie et al., 2014; Hieser et al., 2014; Arnold et al., 2016; Epstein & Smale, 2017; Epstein & Smale, 2018; Kraan, 2017; Epstein et al., 2019a,b; Tidbury, 2020). It originates from Japan but is established currently on the coastlines of New Zealand, Australia, Northern France, Spain, Italy, the UK, Portugal, Belgium, Holland, Argentina, Mexico, and the USA (De Leij et al., 2017). Undaria pinnatifida seems to settle better on artificial substrata (e.g., floats, marinas or piers) than on natural rocky shores among local kelps (Vaz-Pinto et al., 2014) and It is found predominantly in low intertidal to shallow subtidal habitats (Epstein et al., 2019b). Undaria pinnatifida has a wide physiological niche meaning it can occur in both coastal and estuarine environments showing tolerance for varying salinities, turbidity and siltation (Heiser et al., 2014; Epstein & Smale, 2018). Undaria pinnatifida has a greater preference for sites sheltered with low wave exposure and weak tidal streams (Heiser et al., 2014; Epstein & Smale, 2018). In Plymouth Sound (UK), Epstein et al. (2019b) found that within its depth range (+1 to –4 m), Undaria pinnatifida co-existed with seven species of canopy-forming brown macroalgae, including Laminaria hyperborea.  

Undaria pinnatifida was successfully eradicated on a sunken ship in Chatham Islands, New Zealand, by applying a heat treatment of 70°C (Wotton et al., 2004). However, numerous other eradication attempts have failed and as noted by Fletcher & Farrell (1998), once established Undaria pinnatifida resists most attempts at long-term removal. 

Sensitivity assessment. This biotope (IR.HIR.KFaR.Ala.Ldig) is extremely exposed to wave action and found within the sublittoral fringe with moderately strong to weak tidal streams. Sargassum muticum prefers wave sheltered shallow sites in the sublittoral fringe and shallow infralittoral. It was reported to out-compete and replace Saccharina latissima in the Limfjorden and achieve maximum abundance between 1 and 4 m (Staehr et al., 2000; Engelen et al., 2015). However, no evidence of the effects of Sargassum on Alaria esculenta beds was found. Therefore, as Alaria dominates the very exposed coasts it is unlikely that Sargassum will be able to survive within these conditions that characterize the biotope.  

Undaria pinnatifida has the potential to colonize and co-exist in refugia within Laminaria sp. dominated habitats that are within its shallow depth range (+ 1 to – 4 m) and sheltered from wave action. However, like Sargassum muticum, it is highly unlikely that Undaria pinnatifida will be able to colonize or survive within this biotope due to the extreme exposure to wave action that characterizes this biotope. Therefore, resistance to Sargassum or Undaria is assessed as ‘High’, resilience as 'High', and sensitivity is assessed as ‘Not Sensitive’. Overall, confidence is assessed as ‘Low’ due to evidence of variation and the site-specific nature of competition between native kelps and Undaria pinnatifida. 

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

Streblonema sp. is associated with spot disease in kelps and has been found growing on Alaria esculenta (Lein et al. 1991) but no incidence of Alaria esculenta spot disease was found. Corallina officinalis may host several epiphytes of which Titanoderma corallinae is thought to cause tissue damage. Hyperplasia or gall growths are often seen as dark spots on Laminaria digitata and have been associated with endophytic brown filamentous algae. There is no evidence in the literature that infection by microbial pathogens results in the mass death of kelp populations and the kelp themselves are known to regulate bacterial infections through iodine metabolism (Cosse et al., 2009).

Removal of target species

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

Evidence

Alaria esculenta has recently received commercial interest as a consumable product called “Sea Vegetables” or “Atlantic Wakame Kelp”.  However, no studies examining the effect of commercial extraction of Alaria esculenta biotopes were found. Removal of the algal canopy would expose the understorey fauna and flora to increased desiccation. Experimental macroalgal canopy removal experiments conducted in the Isle of Man (Hawkings & Harkin, 1985) found that following removal of the macroalgal canopy the understorey encrusting red algae became bleached and died within a week. Mytilus edulis settlement has also been found significantly higher in close proximity to Alaria esculenta and is thought to increase beneath Alaria esculenta canopies (Bégin et al., 2004).  Therefore, any loss of Alaria esculenta, as a result of commercial extraction, may dramatically affect the understorey community.

Traditionally Laminaria digitata was added to agricultural lands as fertilizers; now Laminaria species are used in a range of different products, with its alginates used in the cosmetic, pharmaceutical and agri-food industries (Kervarec et al., 1999; McHugh, 2003). Laminaria digitata is harvested with a ‘Scoubidou’ (a curved iron hook which is mechanically operated) in France. This device is considered to be selective- only harvesting individuals older than 2 years (Arzel, 2002). France reportedly harvests 75,000t kelp, mainly consisting of Laminaria digitata annually (FAO, 2007). The loss of Laminaria digitata would represent as significant change to IR.HIR.KFaR.Ala.Ldig.

Corallina officinalis is collected for medical purposes; the fronds are dried and converted to hydroxyapatite and used as bone forming material (Ewers et al., 1987). It is also sold as a powder for use in the cosmetic industry. Moderate trampling on articulated coralline algal turf in the New Zealand intertidal (Brown & Taylor, 1999; Schiel & Taylor, 1999) resulted in reduced turf height, declines in turf densities, and loss of crustose bases in some case probably due to loss of the canopy algae and resultant desiccation. Calcification is thought to be an adaptation to grazing and sediment scour (Littler & Kauker 1984). Corallina officinalis produces spores over a protracted period and can colonize artificial substratum within one week in the intertidal (Harlin & Lindbergh 1977; Littler & Kauker, 1984). The crustose base enables Corallina officinalis to survive loss of fronds.

Sensitivity assessment. There is little published evidence for the effects of commercial harvesting of Alaria esculenta. If it is assumed that all canopy forming kelp are removed then resistance would be assessed as ’None‘, resilience would be assessed as ’Medium‘. Sensitivity has been assessed as ’Medium‘. Within IR.HIR.KFaR.AlaAnCrSp and IR.HIR.KFaR.Ala.Myt, monospecific canopies of Alaria esculenta are expected to recover quicker than mixed canopies of Laminaria digitata (as in IR.HIR.KFaR.Ala.Ldig). Sensitivity of the latter would be assessed as follows; resistance ’None‘, resilience as ’High‘, Sensitivity as ’Medium’.

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

Incidental/accidental removal of Alaria esculenta is likely to cause similar effects to that of direct harvesting; hence, the same evidence has been used for both pressure assessments.

Alaria esculenta has recently received commercial interest as a consumable product called “Sea Vegetables” or “Atlantic Wakame Kelp”. However, no studies examining the effect of commercial extraction of Alaria esculenta biotopes were found. Removal of the algal canopy would expose the understorey fauna and flora to increased desiccation. Experimental macroalgal canopy removal experiments conducted in the Isle of Man (Hawkings & Harkin, 1985) found that following the removal of the macroalgal canopy the understorey encrusting red algae became bleached and died within a week. Mytilus edulis settlement has also been found significantly higher in close proximity to Alaria esculenta and is thought to increase beneath Alaria esculenta canopies (Bégin et al., 2004).  Therefore, any loss of Alaria esculenta, as a result of commercial extraction, may dramatically affect the understorey community.

Traditionally Laminaria digitata was added to agricultural lands as fertilizers; now Laminaria species are used in a range of different products, with its alginates used in the cosmetic, pharmaceutical and agri-food industries (Kervarec et al., 1999; McHugh, 2003). Laminaria digitata is harvested with a ‘Scoubidou’ (a curved iron hook which is mechanically operated) in France. This device is considered to be selective- only harvesting individuals older than 2 years (Arzel, 2002). France reportedly harvests 75,000t kelp, mainly consisting of Laminaria digitata annually (FAO, 2007). The loss of Laminaria digitata would represent a significant change to IR.HIR.KFaR.Ala.Ldig.

Corallina officinalis is collected for medical purposes; the fronds are dried and converted to hydroxyapatite and used as bone forming material (Ewers et al., 1987). It is also sold as a powder for use in the cosmetic industry. Moderate trampling on articulated coralline algal turf in the New Zealand intertidal (Brown & Taylor, 1999; Schiel & Taylor, 1999) resulted in reduced turf height, declines in turf densities, and loss of crustose bases in some case probably due to loss of the canopy algae and resultant desiccation. Calcification is thought to be an adaptation to grazing and sediment scour (Littler & Kauker 1984). Corallina officinalis produces spores over a protracted period and can colonize artificial substratum within one week in the intertidal (Harlin & Lindbergh 1977; Littler & Kauker 1984). The crustose base enables Corallina officinalis to survive the loss of fronds.

Sensitivity assessment. There is little evidence for the effects of commercial harvesting of Alaria esculenta. If it is assumed that all canopy-forming kelp are removed then resistance would be assessed as ’None‘, resilience would be assessed as ’Medium‘. Sensitivity has been assessed as ’Medium‘. Within IR.HIR.KFaR.AlaAnCrSp and IR.HIR.KFaR.Ala.Myt, monospecific canopies of Alaria esculenta are expected to recover quicker than mixed canopies of Laminaria digitata (as in IR.HIR.KFaR.Ala.Ldig) Sensitivity of the latter would be assessed as follows; resistance ’None‘, resilience as ’High‘, Sensitivity as ’Medium’.