Saccharina latissima and Gracilaria gracilis with sponges and ascidians on variable salinity infralittoral sediment

Summary

UK and Ireland classification

Description

Shallow kelp community found on stony sediment, in extremely sheltered, variable salinity conditions, with moderately strong tidal currents. The community is characterized by a more sparse covering of Saccharina latissima, particularly when compared to the fully marine version of this sub biotope (SMp.SlatGraFS). Beneath the canopy, the community is characterized by the red algae Gracilaria gracilis, and a variety of faunal species in particular sponges (Suberites ficus and Halichondria panacea) and ascidians (Ascidiella aspersa and Dendrodoa grossularia). The stony substratum provides a surface for attachment for these and many other filter and suspension feeding species, particularly barnacles (Balanus crenatus), hydroids (Hydractinia echinata) and anemones (Urticina felina). Other members of the understorey may include a variety of filamentous and foliose red algae in particular Pterothamnion plumula, and the green alga Ulva. (Information from Connor et al., 2004; JNCC, 2015).

Depth range

0-5 m, 5-10 m

Additional information

-

Listed By

Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS typically occur on a mixture of shallow sediments and rock fractions. The generally sheltered nature of the biotopes allows Saccharina latissima (syn. Laminaria saccharina), Chorda filum, Gracilaria gracilis and other red and brown seaweeds to grow on small stones and shells. Saccharina latissima and Chorda filum are important canopy-forming species and Gracilaria gracilis can characterize the understorey community. The associated community depend on the salinity, with red algae characterizing the full salinity example, while sponges and ascidians characterize the variable salinity example.

In undertaking this assessment of sensitivity, an account is taken of knowledge of the biology of all characterizing species in the biotope. For this sensitivity assessment Saccharina latissima, Chorda filum and Gracilaria gracilis are the primary foci of research, however, it is recognized that the red seaweed species and the abundance of sponges and ascidians also define these biotopes. Examples of important species groups are mentioned where appropriate.

Resilience and recovery rates of habitat

Saccharina latissima (formerly Laminaria saccharina) and Chorda filum are opportunistic seaweeds that have relatively fast growth rates. Saccharina latissima is a perennial kelp that can reach maturity in 15-20 months ((Sjøtun, 1993) and has a life expectancy of 2-4 years (Parke, 1948). Saccharina latissima is widely distributed in the north Atlantic from Svalbard to Portugal (Birket et al., 1998; Conor et al., 2004; Bekby & Moy 2011; Moy & Christie 2012). Chorda filum is widely distributed across the northern hemisphere (Algae Base, 2015). In the North Atlantic, Chorda filum is recorded from Svalbard (Fredriksen et al., 2014) to Northern Portugal (Araújo et al, 2009). Chorda filum is an annual seaweed, completing its life cycle in a single season (Novaczek et al., 1986). Gracilaria gracilis is widely distributed however in the North Atlantic is found from south-west Norway (Rueness, 1977) and extends to South Africa (Anderson et al., 1999). Gracilaria gracilis is widely distributed, in the North Atlantic specifically is found from south-west Norway (Rueness, 2005) and extends to South Africa (Anderson et al., 1999). Gracilaria gracilis is a perennial red seaweed, individuals are composed of an annual erect thallus which grows from a perennial holdfast (Martín et al.,2011).

Saccharina latissima and Chorda filum have heteromorphic life strategies (Edwards, 1998). Mature sporophytes broadcast spawn zoospores from reproductive structures known as sori (South & Burrows, 1967; Birkett et al., 1998b). Zoospores settle onto rock and develop into gametophytes, which following fertilization germinate into juvenile sporophytes. Laminarian zoospores are expected to have a large dispersal range, however, zoospore density and the rate of successful fertilization decreases exponentially with distance from the parental source (Fredriksen et al., 1995). Hence, recruitment can be influenced by the proximity of mature kelp beds producing viable zoospores (Kain, 1979; Fredriksen et al., 1995). Saccharina latissima recruits appear in late winter early spring beyond which is a period of rapid growth, during which sporophytes can reach a total length of 3 m (Werner & Kraan, 2004), in late summer and autumn growth rates slow and spores are released from autumn to winter (Parke, 1948; Lüning, 1979; Birkett et al., 1998b). The overall length of the sporophyte may not change during the growing season due to marginal erosion, but the growth of the blade has been measured at 1.1 cm/day, with a total length addition of ≥2.25 m per year (Birkett et al., 1998b). Chorda filum recruits appear from February (South & Burrows, 1967), beyond which is a period of rapid growth during which sporophytes can reach a length of ≤6 m (South & Burrows, 1967). In culture Chorda filum can reach reproductive maturity and produce zoospores within 186 days (ca 6 months) of settlement, however, the time taken to reach maturity may be locally variable (South & Burrows, 1967). In nature, sporophytes growth slows/stops from October and sporophytes may begin to die off (South & Burrows, 1967; Novaczek et al., 1986).

Saccharina latissima is a rapid colonizing species and appear early in algal succession. For example, Leinaas & Christie (1996) removed Strongylocentrotus droebachiensis from 'Urchin Barrens' and observed a succession effect. Initially, the substrate was colonized by filamentous algae, after a couple of weeks these were outcompeted, and the habitat dominated by Saccharina latissima however this was subsequently outcompeted by Laminaria hyperborea. In the Isle of Man, Kain (1975) cleared sublittoral blocks of Laminaria hyperborea at different times of the year for several years. The first colonizers and succession community differed between blocks and at what time of year the blocks were cleared. Saccharina latissima was an early colonizer, however, within two years of clearance the blocks were dominated by Laminaria hyperborea.

In 2002, a 50.7-83% decline of Saccharina latissima was discovered in the Skaggerak region, South Norway (Moy et al., 2006; Moy & Christie, 2012). Survey results indicated a sustained shift from Saccharina latissima communities to those of ephemeral filamentous algal communities. The reason for the community shift was unknown, but low water movement in wave and tidally sheltered areas combined with the impacts of dense human populations, e.g., increased land run-off, was suggested to be responsible for the dominance of ephemeral turf macro-algae. Multiple stressors such as eutrophication, increasing regional temperature, increased siltation and overfishing may also be acting synergistically to cause the observed habitat shift.

Gracilaria gracilis has a complex life history; reproducing sexually through haploid and diploid spores (Martín et al., 2011) and through vegetative fragmentation (Rueness et al., 1987). Mature individuals consist of erect annual thalli growing from a perennial holdfast (Martín et al., 2011). Vegetative growth is limited to approximately 6 months each year (Kain & Destcombe, 1995) during which thalli can reach 60cm (Bunker et al., 2012). Thalli become reproductively active within two and half months from March-September (Engel & Destombe, 2002). Gracilaria gracilis is recorded throughout the British Isles (AlgaeBase, 2015; NBN, 2015), but SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS core records are restricted to wave and tidally sheltered sites of south-west UK (Connor et al., 2004).

Resilience assessment. Saccharina latissima, Chorda filum and Gracilaria gracilis have the potential to rapidly recover following disturbance. Saccharina latissima has been shown to be an early colonizer within algal succession, appearing within two weeks of clearance, and can reach sexual maturity within 15-20 months. Chorda filum and Gracilaria gracilis have rapid growth rates and are capable of reaching sexual maturity within one year. Resilience has therefore been assessed as ‘High’.

Climate Change Pressures

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ResistanceResilienceSensitivity
Global warming (extreme) [Show more]

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 kelp is strongly influenced by climatic conditions; therefore, kelp species are extremely sensitive to the ongoing ocean warming (Kain, 1979; Van Den Hoek, 1982; Breeman, 1990; Lüning, 1990; Assis et al., 2016; Smale, 2020). Northern distribution boundaries are set by winter temperatures that are lethal, or summer temperatures too low for growth and/or reproduction, while 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 make 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, 2020). 

Saccharina latissima is a polar to temperate macroalgae distributed from Greenland to the coast of Portugal, and in the NW Atlantic is found as far south as New York State, USA. At its southern distribution in New York, temperatures can regularly reach ≥20°C for six weeks or more during summer months (Gerard & Du Bois, 1988). In the UK, sea surface temperatures range between 6-19°C (Huthnance, 2010), and Saccharina latissima is in the middle of its biogeographic range.

Saccharina latissima has an optimal growth temperature between 10-15°C, with growth reducing by 50-70% at 20°C, and all experimental specimens disintegrating after seven days at 23°C (Bolton & Lüning, 1982). The temperature isotherm of 19-20°C has been reported as limiting Saccharina latissima growth (Müller et al., 2009). Temperature is an environmental factor controlling the development of the microscopic stages of Saccharina latissima, with crucial changes in survival, growth, and gametogenesis occurring within a few degrees of its upper thermal limits (Redmond, 2013). The optimal germination temperature for Saccharina latissima is between 2°C and 12°C, with gametophyte survival between 23-25°C (Müller et al., 2009). Germination rates drop at 22°C, with surviving gametophytes smaller than those grown at lower temperatures (Redmond, 2013). Park et al. (2017) observed reductions in the percentage of sporophytes produced at 15°C when compared to values produced at 5°C and 10°C. 

In the field, Saccharina latissima has shown significant regional variation in its acclimation response to changing environmental conditions.  For example, Gerard & Dubois (1988) observed sporophytes of Saccharina latissima that were regularly exposed to ≥20°C tolerated these high temperatures, whereas sporophytes from other populations which rarely experience ≥17°C showed 100% mortality after 3 weeks of exposure to 20°C.

Saccharina latissima has suffered a dramatic decline in the Skagerrak region, Norway, where community structure has shifted from Saccharina latissima forests to communities dominated by filamentous macroalgae (Moy & Christie, 2012). In 2006, Andersen et al. (2011) transplanted Saccharina latissima into areas from where this species had been lost previously to determine whether the kelp could grow and mature. High mortality occurred from August-November each year. In 2008, only six of the seventeen original transplanted Saccharina latissima sporophytes survived (approx. 65% mortality rate). All surviving sporophytes were heavily fouled by epiphytic organisms (estimated cover of 80 & 100%). Between 1960 and 2009, sea surface temperatures in the region had regularly exceeded 20°C and so had the duration at which temperatures remain above 20°C. High sea temperatures have been linked to the slow growth of Saccharina latissima, which is likely due to a decrease in the photosynthetic ability of Saccharina latissima, and an increase in vulnerability to epiphytic loading, bacterial and viral attacks (Anderson et al., 2011).

Assis et al. (2018) predicted that, under the highest emission scenario (RCP 8.5), the range of Saccharina latissima would move northwards, retreating from the coast of Portugal, France and the southwest coast of the UK. The authors projected that, under RCP 2.6, 13% suitable Laminaria hyperborea habitat would be lost from the Western English Channel, while under the RCP 8.5 emission, 87 % of suitable habitat was expected to be lost.

Chorda filum is a cold boreal species, with a wide geographical distribution along the Arctic, Atlantic and Pacific coasts (www.obis.org).  Chorda filum has been reported to have relatively good growth between the temperatures of 5°C and 15°C but reduced or inhibited growth at 20°C (Kawai et al., 2000). Chorda filum has an upper temperature tolerance of 26-28°C (Dieck, 1993). AlthoughLüning (1980) observed that between the temperatures of 15 and 20°C Chorda filum could not reproduce but found that sporophytes could tolerate ≤26 °C. In addition, Lüning (1990) reported gametogenesis to occur at temperatures between 5°C and 10°C in the autumn months. 

Wilson et al. (2015) reported that an increase in sea surface temperature from 1974 to 2010 resulted in biogeographical changes, with declines in abundance of Chorda filum, particularly in the English Channel. Wilson et al. (2015) suggested the declines of Chorda filum could be because the summer temperatures in those southern regions were too high for gametogenesis. 

Gracilaria gracilis has a wide geographical distribution with populations found in the northern and southern hemisphere, from Norway to Argentina and Australia (www.obis.org). In Norway, Gracilaria gracilis exclusively occurs in shallow bays in which summer temperatures exceed 20°C (Rueness, 1977). Biomass production rates vary seasonally, with increased biomass production in the summer months when temperatures are higher and the days are longer (Kain & Destombe, 1995; Martín et al., 2011). Rebello et al. (1996) found the optimal temperature for Gracilaria gracilis growth was 18°C and recorded high growth up to 25.5°C. However, Levy et al. (1990) observed the highest growth rates for Gracilaria gracilis between the temperature of 16°C and 28°C. In addition, Macchiavello et al. (1998) obtained the highest growth rates between 25°C and 28°C and a survival up to 30°C for Gracilaria gracilis.

Ulva sp. are distributed globally (Guiry & Guiry, 2015) and occur in warmer waters than those surrounding the UK suggesting that they can withstand increases in temperature at the pressure benchmark.  Ulva sp. are characteristic of upper shore rock pools, where water and air temperatures are greatly elevated on hot days.  Empirical evidence for thermal tolerance to anthropogenic increases in temperature is provided by the effects of heated effluents on rocky shore communities in Maine, USA.  Ascophyllum and Fucus were eliminated from a rocky shore heated to 27-30 °C by power station effluent whilst Ulva intestinalis (as Enteromorpha intestinalis) increased significantly near the outfall (Vadas et al., 1976).

Sensitivity Assessment. UK populations of Saccharina latissima are found in the middle of the species distribution and are known to be able to survive at higher temperatures than currently experienced around the UK. The ability to tolerate summer seawater temperatures of >20°C in populations at their southern geographic limit is thought to be a genetic adaptation (Gerard & Du Bois, 1988), and maybe crucial in the persistence of this species around the UK, as seawater temperatures rise.

With sea surface temperature around the UK of between 6-19°C (Huthnance, 2010), populations of Saccharina latissima and the understorey community of mixed red seaweeds may be able to adapt to cope with a gradual rise in ocean temperatures of 3°C (middle emission scenario) by the end of this century, leading to maximum summer high temperatures in the south of the UK of 22°C. Gracilaria gracilis is unlikely to be affected by the increase in temperature as the species has been found to grow at temperatures of 25.5°C. However, increasing temperatures are likely to lead to decreased growth and some mortality of Saccharina latissima. Therefore, resistance is assessed as ‘Medium’, and 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 ‘Medium’ sensitivity to ocean warming in the middle emission scenario.

For the high and extreme emission scenario, whereby sea temperatures rise by 4-5°C to potential southern summer temperatures of 23-24°C by the end of this century, Gracilaria gracilis will probably be able to tolerate the rise in temperature, however, Saccharina latissima is likely to be lost from southern England, as gametophytes are not thought to be able to survive at temperatures ≥23°C. This assessment corresponds with the results of ecological niche modelling by Assis et al. (2018), who predicted that Saccharina latissima would be lost from the southwest coast of the UK, because of climate change and the observed reduction in the abundance of Chorda filum from the south coast of the UK (Wilson et al., 2015). Therefore, resistance is assessed as ‘Low’, and resilience is assessed as ‘Very low’. This biotope is assessed as having ‘High’ sensitivity to ocean warming in the high and extreme emission scenarios.

Low
Medium
High
High
Help
Very Low
High
High
High
Help
High
Medium
High
High
Help
Global warming (high) [Show more]

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 kelp is strongly influenced by climatic conditions; therefore, kelp species are extremely sensitive to the ongoing ocean warming (Kain, 1979; Van Den Hoek, 1982; Breeman, 1990; Lüning, 1990; Assis et al., 2016; Smale, 2020). Northern distribution boundaries are set by winter temperatures that are lethal, or summer temperatures too low for growth and/or reproduction, while 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 make 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, 2020). 

Saccharina latissima is a polar to temperate macroalgae distributed from Greenland to the coast of Portugal, and in the NW Atlantic is found as far south as New York State, USA. At its southern distribution in New York, temperatures can regularly reach ≥20°C for six weeks or more during summer months (Gerard & Du Bois, 1988). In the UK, sea surface temperatures range between 6-19°C (Huthnance, 2010), and Saccharina latissima is in the middle of its biogeographic range.

Saccharina latissima has an optimal growth temperature between 10-15°C, with growth reducing by 50-70% at 20°C, and all experimental specimens disintegrating after seven days at 23°C (Bolton & Lüning, 1982). The temperature isotherm of 19-20°C has been reported as limiting Saccharina latissima growth (Müller et al., 2009). Temperature is an environmental factor controlling the development of the microscopic stages of Saccharina latissima, with crucial changes in survival, growth, and gametogenesis occurring within a few degrees of its upper thermal limits (Redmond, 2013). The optimal germination temperature for Saccharina latissima is between 2°C and 12°C, with gametophyte survival between 23-25°C (Müller et al., 2009). Germination rates drop at 22°C, with surviving gametophytes smaller than those grown at lower temperatures (Redmond, 2013). Park et al. (2017) observed reductions in the percentage of sporophytes produced at 15°C when compared to values produced at 5°C and 10°C. 

In the field, Saccharina latissima has shown significant regional variation in its acclimation response to changing environmental conditions.  For example, Gerard & Dubois (1988) observed sporophytes of Saccharina latissima that were regularly exposed to ≥20°C tolerated these high temperatures, whereas sporophytes from other populations which rarely experience ≥17°C showed 100% mortality after 3 weeks of exposure to 20°C.

Saccharina latissima has suffered a dramatic decline in the Skagerrak region, Norway, where community structure has shifted from Saccharina latissima forests to communities dominated by filamentous macroalgae (Moy & Christie, 2012). In 2006, Andersen et al. (2011) transplanted Saccharina latissima into areas from where this species had been lost previously to determine whether the kelp could grow and mature. High mortality occurred from August-November each year. In 2008, only six of the seventeen original transplanted Saccharina latissima sporophytes survived (approx. 65% mortality rate). All surviving sporophytes were heavily fouled by epiphytic organisms (estimated cover of 80 & 100%). Between 1960 and 2009, sea surface temperatures in the region had regularly exceeded 20°C and so had the duration at which temperatures remain above 20°C. High sea temperatures have been linked to the slow growth of Saccharina latissima, which is likely due to a decrease in the photosynthetic ability of Saccharina latissima, and an increase in vulnerability to epiphytic loading, bacterial and viral attacks (Anderson et al., 2011).

Assis et al. (2018) predicted that, under the highest emission scenario (RCP 8.5), the range of Saccharina latissima would move northwards, retreating from the coast of Portugal, France and the southwest coast of the UK. The authors projected that, under RCP 2.6, 13% suitable Laminaria hyperborea habitat would be lost from the Western English Channel, while under the RCP 8.5 emission, 87 % of suitable habitat was expected to be lost.

Chorda filum is a cold boreal species, with a wide geographical distribution along the Arctic, Atlantic and Pacific coasts (www.obis.org).  Chorda filum has been reported to have relatively good growth between the temperatures of 5°C and 15°C but reduced or inhibited growth at 20°C (Kawai et al., 2000). Chorda filum has an upper temperature tolerance of 26-28°C (Dieck, 1993). AlthoughLüning (1980) observed that between the temperatures of 15 and 20°C Chorda filum could not reproduce but found that sporophytes could tolerate ≤26 °C. In addition, Lüning (1990) reported gametogenesis to occur at temperatures between 5°C and 10°C in the autumn months. 

Wilson et al. (2015) reported that an increase in sea surface temperature from 1974 to 2010 resulted in biogeographical changes, with declines in abundance of Chorda filum, particularly in the English Channel. Wilson et al. (2015) suggested the declines of Chorda filum could be because the summer temperatures in those southern regions were too high for gametogenesis. 

Gracilaria gracilis has a wide geographical distribution with populations found in the northern and southern hemisphere, from Norway to Argentina and Australia (www.obis.org). In Norway, Gracilaria gracilis exclusively occurs in shallow bays in which summer temperatures exceed 20°C (Rueness, 1977). Biomass production rates vary seasonally, with increased biomass production in the summer months when temperatures are higher and the days are longer (Kain & Destombe, 1995; Martín et al., 2011). Rebello et al. (1996) found the optimal temperature for Gracilaria gracilis growth was 18°C and recorded high growth up to 25.5°C. However, Levy et al. (1990) observed the highest growth rates for Gracilaria gracilis between the temperature of 16°C and 28°C. In addition, Macchiavello et al. (1998) obtained the highest growth rates between 25°C and 28°C and a survival up to 30°C for Gracilaria gracilis.

Ulva sp. are distributed globally (Guiry & Guiry, 2015) and occur in warmer waters than those surrounding the UK suggesting that they can withstand increases in temperature at the pressure benchmark.  Ulva sp. are characteristic of upper shore rock pools, where water and air temperatures are greatly elevated on hot days.  Empirical evidence for thermal tolerance to anthropogenic increases in temperature is provided by the effects of heated effluents on rocky shore communities in Maine, USA.  Ascophyllum and Fucus were eliminated from a rocky shore heated to 27-30 °C by power station effluent whilst Ulva intestinalis (as Enteromorpha intestinalis) increased significantly near the outfall (Vadas et al., 1976).

Sensitivity Assessment. UK populations of Saccharina latissima are found in the middle of the species distribution and are known to be able to survive at higher temperatures than currently experienced around the UK. The ability to tolerate summer seawater temperatures of >20°C in populations at their southern geographic limit is thought to be a genetic adaptation (Gerard & Du Bois, 1988), and maybe crucial in the persistence of this species around the UK, as seawater temperatures rise.

With sea surface temperature around the UK of between 6-19°C (Huthnance, 2010), populations of Saccharina latissima and the understorey community of mixed red seaweeds may be able to adapt to cope with a gradual rise in ocean temperatures of 3°C (middle emission scenario) by the end of this century, leading to maximum summer high temperatures in the south of the UK of 22°C. Gracilaria gracilis is unlikely to be affected by the increase in temperature as the species has been found to grow at temperatures of 25.5°C. However, increasing temperatures are likely to lead to decreased growth and some mortality of Saccharina latissima. Therefore, resistance is assessed as ‘Medium’, and 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 ‘Medium’ sensitivity to ocean warming in the middle emission scenario.

For the high and extreme emission scenario, whereby sea temperatures rise by 4-5°C to potential southern summer temperatures of 23-24°C by the end of this century, Gracilaria gracilis will probably be able to tolerate the rise in temperature, however, Saccharina latissima is likely to be lost from southern England, as gametophytes are not thought to be able to survive at temperatures ≥23°C. This assessment corresponds with the results of ecological niche modelling by Assis et al. (2018), who predicted that Saccharina latissima would be lost from the southwest coast of the UK, because of climate change and the observed reduction in the abundance of Chorda filum from the south coast of the UK (Wilson et al., 2015). Therefore, resistance is assessed as ‘Low’, and resilience is assessed as ‘Very low’. This biotope is assessed as having ‘High’ sensitivity to ocean warming in the high and extreme emission scenarios.

Low
Medium
High
High
Help
Very Low
High
High
High
Help
High
Medium
High
High
Help
Global warming (middle) [Show more]

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 kelp is strongly influenced by climatic conditions; therefore, kelp species are extremely sensitive to the ongoing ocean warming (Kain, 1979; Van Den Hoek, 1982; Breeman, 1990; Lüning, 1990; Assis et al., 2016; Smale, 2020). Northern distribution boundaries are set by winter temperatures that are lethal, or summer temperatures too low for growth and/or reproduction, while 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 make 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, 2020). 

Saccharina latissima is a polar to temperate macroalgae distributed from Greenland to the coast of Portugal, and in the NW Atlantic is found as far south as New York State, USA. At its southern distribution in New York, temperatures can regularly reach ≥20°C for six weeks or more during summer months (Gerard & Du Bois, 1988). In the UK, sea surface temperatures range between 6-19°C (Huthnance, 2010), and Saccharina latissima is in the middle of its biogeographic range.

Saccharina latissima has an optimal growth temperature between 10-15°C, with growth reducing by 50-70% at 20°C, and all experimental specimens disintegrating after seven days at 23°C (Bolton & Lüning, 1982). The temperature isotherm of 19-20°C has been reported as limiting Saccharina latissima growth (Müller et al., 2009). Temperature is an environmental factor controlling the development of the microscopic stages of Saccharina latissima, with crucial changes in survival, growth, and gametogenesis occurring within a few degrees of its upper thermal limits (Redmond, 2013). The optimal germination temperature for Saccharina latissima is between 2°C and 12°C, with gametophyte survival between 23-25°C (Müller et al., 2009). Germination rates drop at 22°C, with surviving gametophytes smaller than those grown at lower temperatures (Redmond, 2013). Park et al. (2017) observed reductions in the percentage of sporophytes produced at 15°C when compared to values produced at 5°C and 10°C. 

In the field, Saccharina latissima has shown significant regional variation in its acclimation response to changing environmental conditions.  For example, Gerard & Dubois (1988) observed sporophytes of Saccharina latissima that were regularly exposed to ≥20°C tolerated these high temperatures, whereas sporophytes from other populations which rarely experience ≥17°C showed 100% mortality after 3 weeks of exposure to 20°C.

Saccharina latissima has suffered a dramatic decline in the Skagerrak region, Norway, where community structure has shifted from Saccharina latissima forests to communities dominated by filamentous macroalgae (Moy & Christie, 2012). In 2006, Andersen et al. (2011) transplanted Saccharina latissima into areas from where this species had been lost previously to determine whether the kelp could grow and mature. High mortality occurred from August-November each year. In 2008, only six of the seventeen original transplanted Saccharina latissima sporophytes survived (approx. 65% mortality rate). All surviving sporophytes were heavily fouled by epiphytic organisms (estimated cover of 80 & 100%). Between 1960 and 2009, sea surface temperatures in the region had regularly exceeded 20°C and so had the duration at which temperatures remain above 20°C. High sea temperatures have been linked to the slow growth of Saccharina latissima, which is likely due to a decrease in the photosynthetic ability of Saccharina latissima, and an increase in vulnerability to epiphytic loading, bacterial and viral attacks (Anderson et al., 2011).

Assis et al. (2018) predicted that, under the highest emission scenario (RCP 8.5), the range of Saccharina latissima would move northwards, retreating from the coast of Portugal, France and the southwest coast of the UK. The authors projected that, under RCP 2.6, 13% suitable Laminaria hyperborea habitat would be lost from the Western English Channel, while under the RCP 8.5 emission, 87 % of suitable habitat was expected to be lost.

Chorda filum is a cold boreal species, with a wide geographical distribution along the Arctic, Atlantic and Pacific coasts (www.obis.org).  Chorda filum has been reported to have relatively good growth between the temperatures of 5°C and 15°C but reduced or inhibited growth at 20°C (Kawai et al., 2000). Chorda filum has an upper temperature tolerance of 26-28°C (Dieck, 1993). AlthoughLüning (1980) observed that between the temperatures of 15 and 20°C Chorda filum could not reproduce but found that sporophytes could tolerate ≤26 °C. In addition, Lüning (1990) reported gametogenesis to occur at temperatures between 5°C and 10°C in the autumn months. 

Wilson et al. (2015) reported that an increase in sea surface temperature from 1974 to 2010 resulted in biogeographical changes, with declines in abundance of Chorda filum, particularly in the English Channel. Wilson et al. (2015) suggested the declines of Chorda filum could be because the summer temperatures in those southern regions were too high for gametogenesis. 

Gracilaria gracilis has a wide geographical distribution with populations found in the northern and southern hemisphere, from Norway to Argentina and Australia (www.obis.org). In Norway, Gracilaria gracilis exclusively occurs in shallow bays in which summer temperatures exceed 20°C (Rueness, 1977). Biomass production rates vary seasonally, with increased biomass production in the summer months when temperatures are higher and the days are longer (Kain & Destombe, 1995; Martín et al., 2011). Rebello et al. (1996) found the optimal temperature for Gracilaria gracilis growth was 18°C and recorded high growth up to 25.5°C. However, Levy et al. (1990) observed the highest growth rates for Gracilaria gracilis between the temperature of 16°C and 28°C. In addition, Macchiavello et al. (1998) obtained the highest growth rates between 25°C and 28°C and a survival up to 30°C for Gracilaria gracilis.

Ulva sp. are distributed globally (Guiry & Guiry, 2015) and occur in warmer waters than those surrounding the UK suggesting that they can withstand increases in temperature at the pressure benchmark.  Ulva sp. are characteristic of upper shore rock pools, where water and air temperatures are greatly elevated on hot days.  Empirical evidence for thermal tolerance to anthropogenic increases in temperature is provided by the effects of heated effluents on rocky shore communities in Maine, USA.  Ascophyllum and Fucus were eliminated from a rocky shore heated to 27-30 °C by power station effluent whilst Ulva intestinalis (as Enteromorpha intestinalis) increased significantly near the outfall (Vadas et al., 1976).

Sensitivity Assessment. UK populations of Saccharina latissima are found in the middle of the species distribution and are known to be able to survive at higher temperatures than currently experienced around the UK. The ability to tolerate summer seawater temperatures of >20°C in populations at their southern geographic limit is thought to be a genetic adaptation (Gerard & Du Bois, 1988), and maybe crucial in the persistence of this species around the UK, as seawater temperatures rise.

With sea surface temperature around the UK of between 6-19°C (Huthnance, 2010), populations of Saccharina latissima and the understorey community of mixed red seaweeds may be able to adapt to cope with a gradual rise in ocean temperatures of 3°C (middle emission scenario) by the end of this century, leading to maximum summer high temperatures in the south of the UK of 22°C. Gracilaria gracilis is unlikely to be affected by the increase in temperature as the species has been found to grow at temperatures of 25.5°C. However, increasing temperatures are likely to lead to decreased growth and some mortality of Saccharina latissima. Therefore, resistance is assessed as ‘Medium’, and 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 ‘Medium’ sensitivity to ocean warming in the middle emission scenario.

For the high and extreme emission scenario, whereby sea temperatures rise by 4-5°C to potential southern summer temperatures of 23-24°C by the end of this century, Gracilaria gracilis will probably be able to tolerate the rise in temperature, however, Saccharina latissima is likely to be lost from southern England, as gametophytes are not thought to be able to survive at temperatures ≥23°C. This assessment corresponds with the results of ecological niche modelling by Assis et al. (2018), who predicted that Saccharina latissima would be lost from the southwest coast of the UK, because of climate change and the observed reduction in the abundance of Chorda filum from the south coast of the UK (Wilson et al., 2015). Therefore, resistance is assessed as ‘Low’, and resilience is assessed as ‘Very low’. This biotope is assessed as having ‘High’ sensitivity to ocean warming in the high and extreme emission scenarios.

Medium
Medium
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Very Low
High
High
High
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Medium
Medium
High
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Marine heatwaves (high) [Show more]

Marine heatwaves (high)

High emission scenario benchmark: A marine heatwave occurring every two years, with a mean duration of 120 days, and a maximum intensity of 3.5°C. Further detail.

Evidence

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

Saccharina latissima has disappeared almost completely from the Danish estuary Limfjorden, where maximum surface temperatures in summer have increased by 0.7°C per decade over the last 40 years while the number of days with temperatures above 20°C has increased dramatically from 1-2 days year to >25 days year (Pedersen, 2015). Similarly, Saccharina latissima has been lost from the Skagerrak coast of Norway, which is thought to be due to an increase in summer temperatures, coupled with eutrophication (Moy & Christie, 2012).

Under experimental conditions, Nepper-Davidson et al. (2019) exposed a northern (Denmark) population of Saccharina latissima to a simulated three-week heatwave of three different intensities; 18, 21 and 24°C. When exposed to heatwaves of 18 and 21°C there was a decrease in photosynthesis and growth. When a 24°C was simulated, 91% of sporophytes were dead within a week, and the fronds of the few survivors were disintegrating, so the experiment was terminated (Nepper-Davidsen et al., 2019). These results suggest that this species is unlikely to survive heatwaves of the length and magnitude predicted by the end of this century for both the middle and high emission scenarios.

Simonson et al. (2015) investigated the impacts of four temperature treatments (11°C, 14°C, 18°C & 21°C) on Saccharina latissima tissue over three weeksHistological analysis showed temperature mediated tissue damage, including holes, splitting of the medulla, damage to the meristoderm and loss of differentiation between tissue layers at temperatures between 14-21°C. 

Gracilaria gracilis has been reported to have an optimal temperature of 18°C, but high growth rates have been observed up to temperatures of 25.5°C (Rebello et al., 1996). With the highest biomass production rates recorded during the summer months when the temperature is high (Kain & Destombe, 1995; Martín et al., 2011). Therefore, marine heatwaves are unlikely to have an impact on the growth and survival of Gracilaria gracilis. However, no evidence of the effects of marine heatwaves on Gracilaria gracilis was found.

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. A heatwave of this magnitude is unlikely to cause impact to Gracilaria gracilis however the heatwave is likely to cause mass mortality of Saccharina latissima. 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.’ 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. Under this scenario, Saccharina latissima is likely to be already lost from this biotope as a result of rising temperatures (see Global warming) although mortality of any surviving specimens would occur as a result of this projected heatwave. The impacts that a heatwave of this magnitude will have on Gracilaria gracilis is unclear due to the lack of evidence. However, as the main characterizing kelp species is highly likely to be lost, 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.

None
High
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Very Low
High
High
High
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High
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Marine heatwaves (middle) [Show more]

Marine heatwaves (middle)

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

Evidence

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

Saccharina latissima has disappeared almost completely from the Danish estuary Limfjorden, where maximum surface temperatures in summer have increased by 0.7°C per decade over the last 40 years while the number of days with temperatures above 20°C has increased dramatically from 1-2 days year to >25 days year (Pedersen, 2015). Similarly, Saccharina latissima has been lost from the Skagerrak coast of Norway, which is thought to be due to an increase in summer temperatures, coupled with eutrophication (Moy & Christie, 2012).

Under experimental conditions, Nepper-Davidson et al. (2019) exposed a northern (Denmark) population of Saccharina latissima to a simulated three-week heatwave of three different intensities; 18, 21 and 24°C. When exposed to heatwaves of 18 and 21°C there was a decrease in photosynthesis and growth. When a 24°C was simulated, 91% of sporophytes were dead within a week, and the fronds of the few survivors were disintegrating, so the experiment was terminated (Nepper-Davidsen et al., 2019). These results suggest that this species is unlikely to survive heatwaves of the length and magnitude predicted by the end of this century for both the middle and high emission scenarios.

Simonson et al. (2015) investigated the impacts of four temperature treatments (11°C, 14°C, 18°C & 21°C) on Saccharina latissima tissue over three weeksHistological analysis showed temperature mediated tissue damage, including holes, splitting of the medulla, damage to the meristoderm and loss of differentiation between tissue layers at temperatures between 14-21°C. 

Gracilaria gracilis has been reported to have an optimal temperature of 18°C, but high growth rates have been observed up to temperatures of 25.5°C (Rebello et al., 1996). With the highest biomass production rates recorded during the summer months when the temperature is high (Kain & Destombe, 1995; Martín et al., 2011). Therefore, marine heatwaves are unlikely to have an impact on the growth and survival of Gracilaria gracilis. However, no evidence of the effects of marine heatwaves on Gracilaria gracilis was found.

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. A heatwave of this magnitude is unlikely to cause impact to Gracilaria gracilis however the heatwave is likely to cause mass mortality of Saccharina latissima. 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.’ 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. Under this scenario, Saccharina latissima is likely to be already lost from this biotope as a result of rising temperatures (see Global warming) although mortality of any surviving specimens would occur as a result of this projected heatwave. The impacts that a heatwave of this magnitude will have on Gracilaria gracilis is unclear due to the lack of evidence. However, as the main characterizing kelp species is highly likely to be lost, 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.

None
High
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Very Low
High
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High
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Ocean acidification (high) [Show more]

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 emission scenario. Marine autotrophs will generally benefit from ocean acidification, through an increase in the availability of aqueous COfor photosynthesis (Koch et al., 2013). 

Research on most 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).

Under experimental COenrichment at levels expected by the end of this century, germination rates in Saccharina latissima were the same as control samples but gametophyte size increased, suggesting a benefit for juvenile stages of this species (Roleda et al., 2012). Nunes et al. (2015) found that experimental exposure of adult Saccharina latissima to enhanced CO2 led to an increase in net primary production, while Gordillo et al. (2015) found that enhanced CO2 led to increased photosynthesis and growth. In contrast, Iñiguez et al. (2016) found no increase in carbon fixation under elevated CO2 conditions. Although contrasting in findings, these studies show that ocean acidification will not negatively impact Saccharina latissima.

Elevated levels of CO2 have been reported to significantly increase the growth rates of Gracilaria sp. (Gao et al., 1993; Zou & Gao, 2009; Young & Gobler, 2016). Therefore, ocean acidification is not expected to negatively impact Gracilaria sp.

Sensitivity Assessment. Kelp forests live 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). Saccharina latissima and Gracilaria gracilis is not expected to exhibit negative effects from ocean acidification at levels expected for the end of this century. Due to the disturbed nature of the biotope the understorey community can vary locally, therefore impacts to the understorey community has not been included in the assessment. Under both the middle and high emission scenario resistance is assessed as ‘High’, and resilience is assessed as ‘High’ so that sensitivity is assessed as ‘Not sensitive’.

High
High
High
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High
High
High
High
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Not sensitive
High
High
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Ocean acidification (middle) [Show more]

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 emission scenario. Marine autotrophs will generally benefit from ocean acidification, through an increase in the availability of aqueous COfor photosynthesis (Koch et al., 2013). 

Research on most 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).

Under experimental COenrichment at levels expected by the end of this century, germination rates in Saccharina latissima were the same as control samples but gametophyte size increased, suggesting a benefit for juvenile stages of this species (Roleda et al., 2012). Nunes et al. (2015) found that experimental exposure of adult Saccharina latissima to enhanced CO2 led to an increase in net primary production, while Gordillo et al. (2015) found that enhanced CO2 led to increased photosynthesis and growth. In contrast, Iñiguez et al. (2016) found no increase in carbon fixation under elevated CO2 conditions. Although contrasting in findings, these studies show that ocean acidification will not negatively impact Saccharina latissima.

Elevated levels of CO2 have been reported to significantly increase the growth rates of Gracilaria sp. (Gao et al., 1993; Zou & Gao, 2009; Young & Gobler, 2016). Therefore, ocean acidification is not expected to negatively impact Gracilaria sp.

Sensitivity Assessment. Kelp forests live 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). Saccharina latissima and Gracilaria gracilis is not expected to exhibit negative effects from ocean acidification at levels expected for the end of this century. Due to the disturbed nature of the biotope the understorey community can vary locally, therefore impacts to the understorey community has not been included in the assessment. Under both the middle and high emission scenario resistance is assessed as ‘High’, and resilience is assessed as ‘High’ so that sensitivity is assessed as ‘Not sensitive’.

High
High
High
High
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High
High
High
High
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Not sensitive
High
High
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Sea level rise (extreme) [Show more]

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).

Light availability and water turbidity are principle factors in determining kelp depth range (Birkett et al. 1998b), with laminarians being reported to be able to withstand light levels of up to 1% surface irradiance. In Maine, USA, Saccharina latissima is abundant at both turbid and deep sites where light availability averages 2.5% of surface irradiance and have adapted to low-light conditions (Gerard, 1990).

This biotope (SS.SMp.KSwSS.SlatGraFS) occurs on moderately exposed, sheltered, very sheltered and extremely sheltered upper infralittoral and sublittoral fringe zones (JNCC, 2015). Understanding how sea-level rise will affect tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site-dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude. 

Yang et al. (2015) reported Gracilaria gracilis to require high light intensities to maintain normal growth rates. Mensi et al. (2020) recorded the highest daily growth rates from Gracilaria gracilis at depths of <1m and the lowest growth rates in the deeper sample site of >3m. Also, Yang et al. (2015) implied that Gracilaria productivity varies with depth, with increased biomass found in shallower water depths. 

Sensitivity assessment. An increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope, which is already constrained to shallow waters through limits to light availability. The biotope is recorded from 0 to 10 m in depth (JNCC, 2015). 

This biotope may be able to expand its range and migrate landwards to compensate for sea-level rise, if not constrained by lack of tide-swept rock, or human-modified shorelines (IPCC, 2019). If landward migration is not possible, it is expected that depth distribution of this biotope will shrink substantially in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery, due to the increased depth, leading to a reduction in light availability for photosynthesis. 

There is likely to be considerable variation between sites, the relative contribution of wave surge and exposure to habitat suitability, and the depth range occupied by the biotope. Hence, it is difficult to assess the effect of the different sea-level rise scenarios. However, as the biotope can occur from 0-10 m in depth, it is assumed at a sea-level rise of 50 cm, or 70 cm (middle to high emission scenarios) would have limited effect but that a 107 cm rise (the extreme emission scenario) might result in loss of some of the deeper extent of the biotope in some sites. Therefore, resistance is assessed as ‘High’ under the middle and high emission scenarios so that resilience is ‘High’ and sensitivity assessed as ‘Not sensitive’. But resistance may be ‘Medium’ under the extreme emission scenario so that resilience is ‘Very low’ and sensitivity assessed as ‘Medium’, albeit with ‘Low’ confidence.

Medium
Low
NR
NR
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Very Low
High
High
High
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Medium
Low
Low
Low
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Sea level rise (high) [Show more]

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).

Light availability and water turbidity are principle factors in determining kelp depth range (Birkett et al. 1998b), with laminarians being reported to be able to withstand light levels of up to 1% surface irradiance. In Maine, USA, Saccharina latissima is abundant at both turbid and deep sites where light availability averages 2.5% of surface irradiance and have adapted to low-light conditions (Gerard, 1990).

This biotope (SS.SMp.KSwSS.SlatGraFS) occurs on moderately exposed, sheltered, very sheltered and extremely sheltered upper infralittoral and sublittoral fringe zones (JNCC, 2015). Understanding how sea-level rise will affect tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site-dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude. 

Yang et al. (2015) reported Gracilaria gracilis to require high light intensities to maintain normal growth rates. Mensi et al. (2020) recorded the highest daily growth rates from Gracilaria gracilis at depths of <1m and the lowest growth rates in the deeper sample site of >3m. Also, Yang et al. (2015) implied that Gracilaria productivity varies with depth, with increased biomass found in shallower water depths. 

Sensitivity assessment. An increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope, which is already constrained to shallow waters through limits to light availability. The biotope is recorded from 0 to 10 m in depth (JNCC, 2015). 

This biotope may be able to expand its range and migrate landwards to compensate for sea-level rise, if not constrained by lack of tide-swept rock, or human-modified shorelines (IPCC, 2019). If landward migration is not possible, it is expected that depth distribution of this biotope will shrink substantially in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery, due to the increased depth, leading to a reduction in light availability for photosynthesis. 

There is likely to be considerable variation between sites, the relative contribution of wave surge and exposure to habitat suitability, and the depth range occupied by the biotope. Hence, it is difficult to assess the effect of the different sea-level rise scenarios. However, as the biotope can occur from 0-10 m in depth, it is assumed at a sea-level rise of 50 cm, or 70 cm (middle to high emission scenarios) would have limited effect but that a 107 cm rise (the extreme emission scenario) might result in loss of some of the deeper extent of the biotope in some sites. Therefore, resistance is assessed as ‘High’ under the middle and high emission scenarios so that resilience is ‘High’ and sensitivity assessed as ‘Not sensitive’. But resistance may be ‘Medium’ under the extreme emission scenario so that resilience is ‘Very low’ and sensitivity assessed as ‘Medium’, albeit with ‘Low’ confidence.

High
Low
NR
NR
Help
High
High
High
High
Help
Not sensitive
Low
Low
Low
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Sea level rise (middle) [Show more]

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).

Light availability and water turbidity are principle factors in determining kelp depth range (Birkett et al. 1998b), with laminarians being reported to be able to withstand light levels of up to 1% surface irradiance. In Maine, USA, Saccharina latissima is abundant at both turbid and deep sites where light availability averages 2.5% of surface irradiance and have adapted to low-light conditions (Gerard, 1990).

This biotope (SS.SMp.KSwSS.SlatGraFS) occurs on moderately exposed, sheltered, very sheltered and extremely sheltered upper infralittoral and sublittoral fringe zones (JNCC, 2015). Understanding how sea-level rise will affect tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site-dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude. 

Yang et al. (2015) reported Gracilaria gracilis to require high light intensities to maintain normal growth rates. Mensi et al. (2020) recorded the highest daily growth rates from Gracilaria gracilis at depths of <1m and the lowest growth rates in the deeper sample site of >3m. Also, Yang et al. (2015) implied that Gracilaria productivity varies with depth, with increased biomass found in shallower water depths. 

Sensitivity assessment. An increase in sea level height of 50, 70 and 107 cm could have severe repercussions for the extent of this biotope, which is already constrained to shallow waters through limits to light availability. The biotope is recorded from 0 to 10 m in depth (JNCC, 2015). 

This biotope may be able to expand its range and migrate landwards to compensate for sea-level rise, if not constrained by lack of tide-swept rock, or human-modified shorelines (IPCC, 2019). If landward migration is not possible, it is expected that depth distribution of this biotope will shrink substantially in response to a 50, 70 or 107 cm sea-level rise, without the possibility of recovery, due to the increased depth, leading to a reduction in light availability for photosynthesis. 

There is likely to be considerable variation between sites, the relative contribution of wave surge and exposure to habitat suitability, and the depth range occupied by the biotope. Hence, it is difficult to assess the effect of the different sea-level rise scenarios. However, as the biotope can occur from 0-10 m in depth, it is assumed at a sea-level rise of 50 cm, or 70 cm (middle to high emission scenarios) would have limited effect but that a 107 cm rise (the extreme emission scenario) might result in loss of some of the deeper extent of the biotope in some sites. Therefore, resistance is assessed as ‘High’ under the middle and high emission scenarios so that resilience is ‘High’ and sensitivity assessed as ‘Not sensitive’. But resistance may be ‘Medium’ under the extreme emission scenario so that resilience is ‘Very low’ and sensitivity assessed as ‘Medium’, albeit with ‘Low’ confidence.

High
Low
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NR
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High
High
High
High
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Not sensitive
Low
Low
Low
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Hydrological Pressures

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ResistanceResilienceSensitivity
Temperature increase (local) [Show more]

Temperature increase (local)

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

Evidence

The temperature isotherm of 19-20°C has been reported as limiting Saccharina latissima geographic distribution (Müller et al., 2009). Gametophytes can develop in ≤23°C (Lüning, 1990) but the optimal temperature range for sporophyte growth is 10-15°C (Bolton & Lüning, 1982). Bolton & Lüning (1982) experimentally observed that sporophyte growth was inhibited by 50-70% at 20°C and following 7 days at 23°C all specimens completely disintegrated. In the field Saccharina latissima has shown significant regional variation in its acclimation to temperature changes, for example Gerard & Dubois (1988) observed sporophytes of Saccharina latissima which were regularly exposed to ≥20°C could tolerate these temperatures, whereas sporophytes from other populations which rarely experience ≥17°C showed 100% mortality after 3 weeks of exposure to 20°C. Therefore the response of Saccharina latissima to a change in temperatures is likely to be locally variable.

Lüning (1980) observed that Chorda filum could not reproduce at 15-20°C but found that sporophytes could tolerate ≤26°C. The optimal temperature for Gracilaria gracilis growth was found to be 18°C, but high growth was recorded up to 25.5°C (Rebello et al., 1996). Gracilaria gracilis northern range edge is south western Norway where it exclusively occurs in shallow bays in which summer temperatures exceed 20°C (Rueness, 1977).

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 effect of this pressure is likely to be regionally variable

Sensitivity assessment. Ecotypes of Saccharina lattisma have been shown to have different temperature optimums (Dubois, 1988). Acute 5°C increases in temperature for a period of one month combined with high summer temperatures could cause large scale mortality of Saccharina lattisma and inhibit Chorda filum reproduction. Gracilaria gracilis is unlikely to be significantly affected. A 2°C increase in temperature for a period of one year when combined with high summer temperatures could similarly result in large scale mortality of Saccharina lattisima ecotypes. Resistance has been assessed as ‘None’, Resilience as ‘High’. Sensitivity has been assessed as ‘Medium’.

None
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High
High
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Medium
High
High
High
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Temperature decrease (local) [Show more]

Temperature decrease (local)

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

Evidence

Saccharina lattissima and Chorda filum are widespread throughout the arctic. Saccharina lattissima has a lower temperature threshold for sporophyte growth at 0°C (Lüning, 1990). Chorda filum sporophytes can also tolerate 0°C, however at this temperature 99% of zoospores perish (Novaczek et al., (1986). Subtidal red algae can survive at -2°C (Lüning, 1990; Kain & Norton, 1990). The distribution and temperature tolerances of these species suggests they likely be unaffected by temperature decreases assessed within this pressure

Gracilaria gracilis is widespread throughout the UK (Bunker et al., 2012) however has its northern range edge within south west Norway, where it is restricted to shallow bays in which summer temperatures exceed 20°C. Furthermore SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS core records are restricted to the south west UK (Connor et al., 2004), where the average summer Sea Temperature ranges from 12-16°C, and winter 8-13°C (Plymouth: 1981-2010, Beszczynska-Möller & Dye, 2013).

Sensitivity assessment. Considering all the characterizing species are recorded throughout the UK, the reason for the restricted distribution of SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS in the south west UK is unclear however is unlikely due to temperature. SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS has been assessed as ‘Not sensitive’ (resistance and resilience assessed as ‘High’). However due to the proximity of the UK the northern range limit of Gracilaria gracilis and the restricted distribution of SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS in the south west UK confidence has been assessed as ‘Low’.

High
Low
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NR
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High
High
High
High
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Not sensitive
Low
Low
Low
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Salinity increase (local) [Show more]

Salinity increase (local)

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

Evidence

Karsten (2007) tested the photosynthetic ability of Saccharina latissima under acute 2 and 5 day exposure to salinity treatments ranging from 5-60 psu. A control experiment was also carried at 34 psu . Saccharina latissima showed high photosynthetic ability at >80% of the control levels between 25-55 psu.

Optimal salinities for Gracilaria gracilis growth have been recorded at 30‰ (Rebello et al., 1996), however Gracilaria gracilis can reportedly tolerate wide salinity fluctuations (Bunker et al., 2012).  Furthermore, Chorda filum and Gracilaria gracilis can be found in rock pools where salinity is likely to be highly variable (South & Burrows, 1967; Engel & Destombe, 2002). High air temperatures and direct sunlight can cause high surface water evaporation, and resultant increases in salinity within the rock pool. The extent of temperature and salinity change is affected by the frequency and time of day at which tidal inundation occurs, and the size of the pool (Pyefinch, 1943). It should be noted however that local populations may be acclimated to the prevailing salinity regime and may therefore exhibit different tolerances to other populations subject to different salinity conditions and therefore caution should be used when inferring tolerances. It is therefore likely that Chorda filum and Gracilaria gracilis may be tolerant of short-term hyper (≥40‰) and hypo saline conditions. Other members of the community, e.g Ceramium sp. and Ulva are found widely across the intertidal, while the sponges and ascidians probably vary in hypersaline tolerance, although no evidence was available. The effects of long-term exposure to hypersaline conditions are unknown.

Sensitivity assessment. The evidence suggests that Saccharina latissima, Chorda filum and Gracilaria gracilis may tolerate short-term exposure to hypersaline conditions (≥40‰). An increase in salinity to ≥40‰ may however be above the optima for a proportion of the characterizing species and cause a decline in growth. Resistance has been assessed as ‘Medium’, resilience as ‘High’. The sensitivity of this biotope to an increase in salinity has been assessed as ‘Low’.

Medium
Low
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High
High
High
High
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Low
Low
Low
Low
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Salinity decrease (local) [Show more]

Salinity decrease (local)

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

Evidence

Karsten (2007) tested the photosynthetic ability of Saccharina latissima under acute 2 and 5 day exposure to salinity treatments ranging from 5-60 psu. A control experiment was also carried at 34 psu. Saccharina latissima showed high photosynthetic ability at >80% of the control levels between 25-55 psu. Hyposaline treatment of 10-20 psu led to a gradual decline of photosynthetic ability. After 2 days at 5 psu Saccharina latissima showed a significant decline in photosynthetic ability at approx. 30% of control. After 5 days at 5 psu Saccharina latissima specimens became bleached and showed signs of severe damage. The experiment was conducted on Saccharina latissima from the Arctic, and the authors suggest that at extremely low water temperatures (1-5°C) macroalgae acclimation to rapid salinity changes could be slower than at temperate latitudes. It is therefore possible that resident Saccharina latissima of the UK maybe be able to acclimate to salinity changes more effectively.

Chorda filum is tolerant of low salinities (Wilce, 1959; Hayren, I940; Norton & South, 1969), and has been recorded at Björnholm, Finland at a salinity as low as 5.15‰ (Hayren, I940). Norton & South (1969) observed that Chorda filum could develop sporophytes at ≥5‰ under laboratory conditions, however at low salinities the time taken to develop into sporophytes took 65 days at 5‰, or 16 days at 35‰. It was also noted that below 9%o sporophytes did not grow above 2 mm in length.

Optimal salinities for Gracilaria gracilis growth have been recorded at 30‰ (Rebello et al., 1996), however Gracilaria gracilis can reportedly tolerate wide salinity fluctuations (Bunker et al., 2012).  Furthermore, Chorda filum and Gracilaria gracilis can be found in rock pools where salinity is likely to be highly variable (South & Burrows, 1967; Engel & Destombe, 2002). High air temperatures and direct sunlight can cause high surface water evaporation, and resultant increases in salinity within the rock pool. The extent of temperature and salinity change is affected by the frequency and time of day at which tidal inundation occurs, and the size of the pool (Pyefinch, 1943). It should be noted however that local populations may be acclimated to the prevailing salinity regime and may therefore exhibit different tolerances to other populations subject to different salinity conditions and therefore caution should be used when inferring tolerances. It is therefore likely that Chorda filum and Gracilaria gracilis may be tolerant of short-term hyper (≥40‰) and hypo saline conditions. Other members of the community, e.g. Ceramium sp. and Ulva are found widely across the intertidal, while the sponges and ascidians probably vary in hypersaline tolerance, although no evidence was available. The effects of long-term exposure to hypersaline conditions are unknown.

Sensitivity assessment. SlatGraFS is record at ‘Full’ salinity while SlatGraVS occur at Full and Variable (18-40 psu). A decrease in one MNCR category (i.e. from Full to Variable) would probably result in a change in SlatGraFS to SlatGraVS, although SlatGRaVS would not be affected. However, in variable salinity examples of the SlatGraVS biotope, a reduction on one MNCR category from variable to reduced (18 -30 psu) would probably remain within the tolerance range of the characteristic species, most of which are found in the intertidal exposed to short-term freshwater runoff or occur in estuaries. However, there may be a loss of species richness. Resistance has been assessed as ‘Medium’ resilience as ‘High’. Sensitivity of this biotope to a decrease in salinity has been assessed as ‘Low’.

Medium
High
High
High
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High
High
Medium
High
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Low
High
High
High
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Water flow (tidal current) changes (local) [Show more]

Water flow (tidal current) changes (local)

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

Evidence

Peteiro & Freire (2013) measured Saccharina latissima growth from 2 sites, the 1st had maximal water velocities of 0.3 m/sec and the 2nd 0.1 m/sec. At site 1 Saccharina latissima had significantly larger biomass than at site 2 (16 kg/m to 12 kg/m respectively). Peteiro & Freire (2013) suggested that faster water velocities were beneficial to Saccharina latissima growth. However, Gerard & Mann (1979) measured Saccharina latissima productivity at greater water velocities and found Saccharina latissima productivity is reduced in moderately strong tidal streams (≤1 m/sec) when compared to weak tidal streams (<0.5 m/sec).

Chorda filum sporophytes often grow on unstable objects, such as pebbles and shell. Owing to the typically unstable substratum which Chorda filum grows on, whole populations can be moved during storms and deposited in more sheltered locations where development will continue (South & Burrows, 1967). The survival of Chorda filum sporophytes following transport of their attached substrata indicates the species is relatively tolerant to changes in water flow or wave action.

SS.SMp.KSwSS.SlatGraFS and SS.SMp.KSwSS.SlatGraVS are found in a range of tidal streams (<0.5-1.5m/sec-weak) (Connor et al., 2004) indicating that Gracilaria gracilis is tolerant to high water movement. The mobile nature of the sediment within this biotope allows opportunistic Laminarians to dominate the canopy, and inhibits significant colonisation by other kelps. The range of tidal streams in which these biotopes are found suggests that an increase in tidal streams of 0.1-0.2m/sec would not have a significant effect on the biotope

Sensitivity assessment. A change of 0.1 m/s to 0.2 m/s is not likely to dramatically affect biotope structure. Resistance has been assessed as ‘High’, resilience as ‘High’. Sensitivity has been assessed as ‘Not Sensitive’. 

High
High
High
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High
High
High
High
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Not sensitive
High
High
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Emergence regime changes [Show more]

Emergence regime changes

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

Evidence

The biotopes (SlatGraFS and SlatGraVS) are shallow sublittoral biotopes recorded from 0-10m BCD, and as such could likely be exposed during some extreme low tides. Saccharina latissima, Chorda filum and Gracilaria gracilis can grow in the sub-littoral fringe and/or intertidal (South & Burrows, 1967; Engel & Destombe, 2002; White & Marshall, 2007).

An increase in emergence will result in an increased risk of desiccation and mortality of Saccharina latissima, Chorda filum and Gracilaria gracilis. Removal of macro-algae canopy may also increase desiccation and mortality of the undergrowth red seaweed community (Hawkins & Harkin, 1985). Providing that suitable substrata are present, the biotope is likely to re-establish further down the shore within a similar emergence regime to that which existed previously. Similarly, a decrease in emergence could allow the biotope to extend up the shore (depending on substratum availability) although the its lower extent may be lost.

Sensitivity assessment. Resistance has been assessed as ‘Low’. Resilience as ‘High’. The sensitivity of this biotope to a change in emergence is considered as ‘Low’.

Low
Medium
High
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High
High
Low
High
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Low
Medium
Low
High
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Wave exposure changes (local) [Show more]

Wave exposure changes (local)

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

Evidence

Birkett et al. (1998) suggested that Saccharina latissima is rarely present in areas of wave exposure, where it is out-competed by Laminaria hyperborea. However The mobile nature of the sediment within this biotope allows opportunistic Laminarians, such as Saccharina latissima, to dominate the canopy, and inhibits significant colonisation by other kelps. Chorda filum sporophytes often grow on unstable objects, such as pebbles and shell. Owing to the typically unstable substratum which Chorda filum grows on, whole populations can be moved during storms and deposited in more sheltered locations where development will continue (South & Burrows, 1967). The survival of Chorda filum sporophytes following transport of their attached substrata indicates the species is relatively tolerant to changes in water flow or wave action.

Bunker et al. (2012) reports that Gracilaria gracilis is most common in wave sheltered sites. However, SS.SMp.KSwSS.SlatGraFS and SS.SMp.KSwSS.SlatGraVS are recorded from moderately exposed to extremely sheltered sites(Connor et al., 2004).

Sensitivity assessment. An increase in local wave height (e.g. to strong or moderately strong exposure) may increase local sediment mobility, potentially increase dislodgment or relocation of the characterizing species (South & Burrows, 1967; Birkett et al., 1998). An increase in wave exposure, may therefore result in significant change to or loss of the biotope. However, an increase in nearshore significant wave height of 3-5% is not likely to have a significant effect on biotope structure. Resistance has been assessed as ‘High’, Resilience as ‘High’. Sensitivity has been assessed as ‘Not Sensitive’ at the benchmark level.

High
Medium
High
High
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High
High
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Not sensitive
Medium
High
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Chemical Pressures

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ResistanceResilienceSensitivity
Transition elements & organo-metal contamination [Show more]

Transition elements & organo-metal contamination

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

Evidence

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

Bryan (1984) suggested that the general order for heavy metal toxicity in seaweeds is: Organic Hg > inorganic Hg > Cu > Ag > Zn > Cd > Pb. Cole et al., (1999) reported that Hg was very toxic to macrophytes. Similarly, Hopkin & Kain (1978) demonstrated sub-lethal effects of heavy metals on kelp gametophytes and sporophytes, including reduced growth and respiration. Sheppard et al. (1980) noted that increasing levels of heavy metal contamination along the west coast of Britain reduced species number and richness in holdfast fauna, except for suspension feeders which became increasingly dominant. Gastropods may be relatively tolerant of heavy metal pollution (Bryan, 1984). Although macroalgae species may not be killed, except by high levels of contamination, reduced growth rates may impair the ability of the biotope to recover from other environmental disturbances. Thompson & Burrows (1984) observed the growth of Saccharina latissima sporophyte growth was significantly inhibited at 50 µg Cu /l, 1000 µg Zn/l and 50 µg Hg/l. Zoospores were found to be more intolerant and significant reductions in survival rates were observed at 25 µg Cu/l, 1000 µg Zn/l and 5 µg/l.

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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Hydrocarbon & PAH contamination [Show more]

Hydrocarbon & PAH contamination

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

Evidence

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

The mucilaginous slime layer coating of Laminarians may protect them from smothering by oil. Hydrocarbons in solution reduce photosynthesis and may be algicidal. However, Holt et al. (1995) reported that oil spills in the USA and from the Torrey Canyon had little effect on kelps. Similarly, surveys of subtidal communities at a number sites between 1-22.5 m below chart datum showed no noticeable impacts of the Sea Empress oil spill and clean up (Rostron & Bunker, 1997) or during experimental release of untreated oil in Baffin Island, Canada (Cross et al., 1987). Laboratory studies of the effects of oil and dispersants on several red algae species (Grandy 1984) concluded that they were all sensitive to oil/ dispersant mixtures, with little differences between adults, sporelings, diploid or haploid life stages.

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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Synthetic compound contamination [Show more]

Synthetic compound contamination

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

Evidence

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

O'Brian & Dixon (1976) suggested that red algae were the most sensitive group of macrophytes to oil and dispersant contamination (see Smith, 1968). Saccharina latissima has also been found to be sensitive to antifouling compounds. Johansson (2009) exposed samples of Saccharina latissima to several antifouing compounds, observing chlorothalonil, DCOIT, dichlofluanid and tolylfluanid inhibited photosynthesis. Exposure to Chlorothalonil and tolylfluanid, was also found to continue inhibiting oxygen evolution after exposure had finished, and may cause irreversible damage.

Smith (1968) observed that epiphytic and benthic red algae were intolerant of dispersant or oil contamination during the Torrey Canyon oil spill; only the epiphytes Crytopleura ramosa and Spermothamnion repens and some tufts of Jania rubens survived together with Osmundea pinnatifida, Gigartina pistillata and Phyllophora crispa from the sublittoral fringe.

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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Radionuclide contamination [Show more]

Radionuclide contamination

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

Evidence

No evidence

Not relevant (NR)
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Not relevant (NR)
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No evidence (NEv)
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Introduction of other substances [Show more]

Introduction of other substances

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

Evidence

This pressure is Not assessed.

Not Assessed (NA)
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Not assessed (NA)
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Not assessed (NA)
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De-oxygenation [Show more]

De-oxygenation

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

Evidence

Reduced oxygen concentrations can 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 environmental conditions are transient. If levels do drop below 4 mg/l negative effects on these organisms can be expected with adverse effects occurring below 2 mg/l (Cole et al., 1999).

Sensitivity assessment. Reduced oxygen levels are likely to inhibit photosynthesis and respiration but not cause a loss of the macroalgae population directly. Resistance has been assessed as ‘High’ and resilience as ‘High’. Sensitivity has been assessed as ‘Not sensitive’ at the benchmark level.

High
Medium
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High
Medium
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Not sensitive
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Nutrient enrichment [Show more]

Nutrient enrichment

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

Evidence

Conolly & Drew (1985) found Saccharina latissima sporophytes had relatively higher growth rates when in close proximity to a sewage outlet in St Andrews, UK, compared to other sites along the east coast of Scotland. At St Andrews nitrate levels were 20.22µM, which represents an approx. 25% increase compared to other sites (approx. 15.87 µM). Handå et al. (2013) also reported Saccharina latissima sporophytes grew approx. 1% faster per day when in close proximity to Norwegian salmon farms, where elevated ammonium could be readily absorbed by sporophytes. Read et al. (1983) reported after the installation of a new sewage treatment works, which reduced the suspended solid content of liquid effluent by 60% in the Firth of Forth, Saccharina latissima became abundant where previously it had been absent. Bokn et al. (2003) conducted a nutrient loading experiment on intertidal fucoids. Within 3 years of the experiment no significant effect was observed in the communities, however 4-5 years into the experiment a shift occurred from perennials to ephemeral algae. Although Bokn et al. (2003) focussed on fucoids the results could indicate that long-term (>4 years) nutrient loading can result in community shift to ephemeral algae species. Disparities between the findings of the aforementioned studies are likely to be related to the level of organic enrichment. Smit (2002) suggested that dissolved inorganic nitrogen from fish factory waste in Saldanha Bay, South Africa could maintain Gracilaria gracilis growth when natural nutrient sources were low. 

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types. Johnston & Roberts (2009) however also highlighted that macroalgal communities are relative tolerant to contamination, but that contaminated communities can have low diversity assemblages which are dominated by opportunistic and fast growing species (Johnston & Roberts, 2009 and references therein).

Sensitivity assessment. Although short-term exposure (<4 years) to nutrient enrichment may not affect seaweeds directly, indirect effects such as turbidity may significantly affect photosynthesis and result in reduced growth and reproduction and increased competition form fast growing but ephemeral species. However, this biotope is considered to be 'Not sensitive' at the pressure benchmark, that assumes compliance with good status as defined by the WFD.

Not relevant (NR)
NR
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Not relevant (NR)
NR
NR
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Not sensitive
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Organic enrichment [Show more]

Organic enrichment

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

Evidence

Conolly & Drew (1985) found Saccharina latissima sporophytes had relatively higher growth rates when in close proximity to a sewage outlet in St Andrews, UK, compared to other sites along the east coast of Scotland. At St Andrews nitrate levels were 20.22µM, which represents an approx. 25% increase compared to other sites (approx. 15.87 µM). Handå et al. (2013) also reported Saccharina latissima sporophytes grew approx. 1% faster per day when in close proximity to Norwegian salmon farms, where elevated ammonium could be readily absorbed by sporophytes. Read et al. (1983) reported after the installation of a new sewage treatment works, which reduced the suspended solid content of liquid effluent by 60% in the Firth of Forth, Saccharina latissima became abundant where previously it had been absent. Bokn et al. (2003) conducted a nutrient loading experiment on intertidal fucoids. Within 3 years of the experiment no significant effect was observed in the communities, however 4-5 years into the experiment a shift occurred from perennials to ephemeral algae. Although Bokn et al. (2003) focussed on fucoids the results could indicate that long-term (>4 years) nutrient loading can result in community shift to ephemeral algae species. Disparities between the findings of the aforementioned studies are likely to be related to the level of organic enrichment.

Johnston & Roberts (2009) conducted a meta-analysis, which reviewed 216 papers to assess how a variety of contaminants (including sewage and nutrient loading) affected 6 marine habitats (including subtidal reefs). A 30-50% reduction in species diversity and richness was identified from all habitats exposed to the contaminant types. Johnston & Roberts (2009) however also highlighted that macroalgal communities are relatively tolerant to contamination, but that contaminated communities can have low diversity assemblages which are dominated by opportunistic and fast growing species (Johnston & Roberts, 2009 and references therein). Organic enrichment may also result in phytoplankton blooms that increase turbidity and therefore may negatively impact photosynthesis.

Sensitivity assessment. Although short-term exposure (<4 years) to organic enrichment may not affect seaweeds directly, indirect effects such as turbidity may significantly affect photosynthesis and result in reduced growth and reproduction and increased competition from fast growing but ephemeral species. Resistance has been assessed as ‘Medium’, resilience as ‘High’. Sensitivity has been assessed as ’Low’.

Medium
Medium
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High
High
Medium
High
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Low
Medium
Medium
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Physical Pressures

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ResistanceResilienceSensitivity
Physical loss (to land or freshwater habitat) [Show more]

Physical loss (to land or freshwater habitat)

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

Evidence

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

None
High
High
High
Help
Very Low
High
High
High
Help
High
High
High
High
Help
Physical change (to another seabed type) [Show more]

Physical change (to another seabed type)

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

Evidence

If sediment were replaced with rock or artificial substrata, this would represent a fundamental change to the biotope (Macleod et al., 2014). All the characterizing species within this biotope can grow in rock biotopes (Birkett et al., 1998; Connor et al., 2004), however SS.SMp.KSwSS are, by definition, sediment biotopes and introduction of rock would change them into a rock based habitat complex.

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

None
High
High
High
Help
Very Low
High
High
High
Help
High
High
High
High
Help
Physical change (to another sediment type) [Show more]

Physical change (to another sediment type)

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

Evidence

The benchmark for this pressure refers to a change in one Folk class. The pressure benchmark originally developed by Tillin et al., (2010) used the modified Folk triangle developed by Long (2006) which simplified sediment types into four categories: mud and sandy mud, sand and muddy sand, mixed sediments and coarse sediments. The change referred to is therefore a change in sediment classification rather than a change in the finer-scale original Folk categories (Folk, 1954). The change in one Folk class is considered to relate to a change in classification to adjacent categories in the modified Folk triangle. For mixed sediments and sand and muddy sand habitats a change in one Folk class may refer to a change to any of the sediment categories. Dredging and dumping of sediment, and infrastructure developments, can lead to changes in sediment character.

SS.SMp.KSwSS.SlatGraFS & SS.SMp.KSwSS.SlatGraVS occur on mixed substrata, therefore within this pressure a change in one folk class relates to a change to either “Coarse sediment”, “Mud and sandy Mud” and “Sand and sandy mud”. Macroalgae are likely to successfully recruit onto the larger sediment/small rock fractions within these biotopes (e.g. gravel, pebbles, cobbles).Therefore, if the proportion of stabilised large sediment/small rock fractions increased this may benefit these biotopes. Conversely if the proportion of smaller sediment fractions increased within these biotopes (as with “Mud and sandy Mud” and “Sand and sandy mud”) then macro-algal recruitment would likely be significantly reduced.

Sensitivity assessment. Resistance has been assessed as ‘None’, resilience as Very low (the pressure is a permanent change), and sensitivity as High. 

None
Low
NR
NR
Help
Very Low
High
High
High
Help
High
Low
Low
Low
Help
Habitat structure changes - removal of substratum (extraction) [Show more]

Habitat structure changes - removal of substratum (extraction)

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

Evidence

SS.SMp.KSwSS.SlatGraFS and SS.SMp.KSwSS.SlatGraVS are found on a varied mixture of sediment and rock fractions. Extraction of substratum to 30 cm is likely to remove small sediment fractions (e.g. gravel, cobbles) and may mobilize the remaining larger rock fractions (e.g. boulders) causing high mortality within the resident community. All characterizing species have rapid growth rates and are likely to recover within 2 years.

Sensitivity assessment. Resistance has been assessed as ‘None’, Resilience as ‘High’. Sensitivity has been assessed as ‘Medium’. 

None
High
High
High
Help
High
High
High
High
Help
Medium
High
High
High
Help
Abrasion / disturbance of the surface of the substratum or seabed [Show more]

Abrasion / disturbance of the surface of the substratum or seabed

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

Evidence

Abrasion of the substratum e.g. from bottom or pot fishing gear, cable laying etc. may cause localised mobility of the substrata and mortality of the resident community. The effect would be situation dependent however if bottom fishing gear were towed over a site it may mobilise a high proportion of the substrata and cause high mortality in the resident community.

The effect of trampling on shallow algal communities was examined by a single Mediterranean study (Milazzo et al., 2002). Experimental trampling of 18 transects were carried out at 0, 10, 25, 50, 100 and 150 passes and the community examined immediately after and three months later in the shallow infralittoral (0.3-0.5 m below mean low water). Percentage cover and canopy were significantly affected by trampling, the degree of effect increasing in proportion with trampling intensity. Intermediate trampling treatments (25, 50 and 100 tramples) were similar in effect but significantly different from 0 and 10 tramples. After 150 tramples, percentage cover was significantly lower. Erect macroalgae were particularly susceptible, e.g. the canopy forming Cystoseira brachicarpa v. balearica and Dictyota mediterranea. At low to intermediate trampling intensity, Dictyota mediterranea was strongly damaged while Cystoseira brachicarpa v. balearica lost fronds. At high trampling intensities, D. mediterranea was completely removed while C. brachicarpa v. balearica was reduced to holdfasts. Low to intermediate trampling intensities (10, 25, 50 tramples) resulted in a loss of algal biomass of 50 g/m2 , while 100 or 150 tramples resulted in a loss of ca 150 g/m2 . Recovery was incomplete after three months and significant differences in effect were still apparent between trampling treatments. Overall, trampling reduced percentage algal cover and canopy. However, the study focused on the canopy forming species and lower turf forming species were not mentioned. In summary the above evidence suggests that shallow infralittoral algal communities are susceptible to the effects of trampling by pedestrians. Again the canopy forming, erect species seem to be the most susceptible. Trampling of sublittoral fringe communities could occur as coasteerers haul themselves out of the water at the bottom of the shore. Therefore, sublittoral fringe communities in the UK could be susceptible but there is limited evidence at present (Tyler-Walters 2005).

Sensitivity assessment. Resistance has been assessed as ‘None’, Resilience as ‘High’. Sensitivity has been assessed as ‘Medium’.

None
Low
NR
NR
Help
High
High
High
High
Help
Medium
Low
Low
Low
Help
Penetration or disturbance of the substratum subsurface [Show more]

Penetration or disturbance of the substratum subsurface

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

Evidence

Penetration and/or disturbance of the substrate below the surface of the seabed, may cause localised mobility of the substrata, remove kelps and bury read algae, remove large rock fractions (e.g. cobble and pebbles), and remove a proportion of the infuana and result in and mortality of the resident community. However, there is no direct evidence of the effects of mobile gear (e.g. scallop dredges) on these habitats.

Sensitivity assessment. Resistance has been assessed as ‘None’, Resilience as ‘High’. Sensitivity has been assessed as ‘Medium’.

None
Low
NR
NR
Help
High
High
High
High
Help
Medium
Low
Low
Low
Help
Changes in suspended solids (water clarity) [Show more]

Changes in suspended solids (water clarity)

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

Evidence

Suspended Particle Matter (SPM) concentration has a positive linear relationship with sub surface light attenuation (Kd) (Devlin et al., 2008). Light availability and water turbidity are principal factors in determining depth range at which macro-algae can be found (Birket et al., 1998). Light penetration influences the maximum depth at which laminarians 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 100m in the Mediterranean to only 6-7m in the silt laden German Bight. In Atlantic European waters, the depth limit is typically 35m. In very turbid waters the depth at which kelp is found may be reduced, or in some cases excluded completely (e.g. Severn Estuary), because of the alteration in light attenuation by suspended sediment (Lüning, 1990; Birkett et al. 1998). Laminarians show a decrease of 50% photosynthetic activity when turbidity increases by 0.1/m (light attenuation coefficient =0.1-0.2/m; Staehr & Wernberg, 2009).

Sensitivity Assessment. An increase in water clarity from clear to intermediate (10-100mg/l) represent a change in light attenuation of ca 0.67-6.7 Kd/m, and is likely to result in a greater than 50% reduction in photosynthesis of Laminaria spp. Therefore the dominant kelp species will probably suffer a severe decline and resistance to this pressure is assessed as ‘None’.  Resilience is probably to this pressure is defined as ‘Medium’ at the benchmark. Hence, this biotope is regarded as having a sensitivity of ‘Medium ‘to this pressure.

None
High
High
High
Help
High
High
High
High
Help
Medium
High
High
High
Help
Smothering and siltation rate changes (light) [Show more]

Smothering and siltation rate changes (light)

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

Evidence

Smothering by sediment e.g. 5 cm material during a discrete event, is unlikely to damage mature examples of Saccharina latissima, Chorda filum or Gracilaria gracilis but may provide a physical barrier to zoospore settlement and therefore could negatively impact on recruitment processes (Moy & Christie, 2012). Laboratory studies showed that kelp and gametophytes can survive in darkness for between 6-16 months at 8 deg C and would probably survive smothering by a discrete event and once returned to normal conditions gametophytes resumed growth or maturation within 1 month (Dieck, 1993).

SS.SMp.KSwSS biotopes are all recorded from moderately strong to weak tidal streams (≤0.5-1.5 m/sec) (Connor et al., 2004). In tidally exposed biotopes deposited sediment is unlikely to remain for more than a few tidal cycles (due to water flow or wave action). In sheltered biotopes deposited sediment could remain but are unlikely to remain for longer than a year.

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

High
Low
NR
NR
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High
High
High
High
Help
Not sensitive
Low
Low
Low
Help
Smothering and siltation rate changes (heavy) [Show more]

Smothering and siltation rate changes (heavy)

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

Evidence

Smothering by sediment e.g. 30 cm material during a discrete event, is unlikely to damage mature examples of Saccharina latissima, Chorda filum or Gracilaria gracilis but may provide a physical barrier to zoospore settlement and therefore could negatively impact on recruitment processes (Moy & Christie, 2012). Laboratory studies showed that kelp and gametophytes can survive in darkness for between 6-16 months at 8deg C and would probably survive smothering by a discrete event and once returned to normal conditions gametophytes resumed growth or maturation within 1 month (Dieck, 1993).

SS.SMp.KSwSS biotopes are all recorded from moderately strong to weak tidal streams (≤0.5-1.5 m/sec) (Connor et al., 2004). In tidally exposed biotopes deposited sediment is unlikely to remain for more than a few tidal cycles (due to water flow or wave action). In sheltered biotopes deposited sediment could remain but are unlikely to remain for longer than a year.

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

Medium
Low
NR
NR
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High
High
High
High
Help
Low
Low
Low
Low
Help
Litter [Show more]

Litter

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

Evidence

Not assessed.

Not Assessed (NA)
NR
NR
NR
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Not assessed (NA)
NR
NR
NR
Help
Not assessed (NA)
NR
NR
NR
Help
Electromagnetic changes [Show more]

Electromagnetic changes

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

Evidence

No evidence

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
No evidence (NEv)
NR
NR
NR
Help
Underwater noise changes [Show more]

Underwater noise changes

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

Evidence

Not relevant

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

Introduction of light or shading

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

Evidence

There is no evidence to suggest that anthropogenic light sources would affect macro-algae. Shading of the biotope (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 seaweed abundance.

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

Low
Low
NR
NR
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High
Low
NR
NR
Help
Low
Low
Low
Low
Help
Barrier to species movement [Show more]

Barrier to species movement

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

Evidence

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.

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Death or injury by collision [Show more]

Death or injury by collision

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

Evidence

Not relevant. Collision from grounding vessels is addressed under abrasion pressure.

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
Visual disturbance [Show more]

Visual disturbance

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

Evidence

Not relevant

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

Biological Pressures

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ResistanceResilienceSensitivity
Genetic modification & translocation of indigenous species [Show more]

Genetic modification & translocation of indigenous species

Benchmark. Translocation of indigenous species or the introduction of genetically modified or genetically different populations of indigenous species that may result in changes in the genetic structure of local populations, hybridization, or change in community structure. Further detail

Evidence

There is no evidence for translocation of Saccharina latissima, Chorda filum or Gracilaria gracilis over significant geographic distances. Gracilaria gracilis is a commercial species valued for agar production (Marinho-Soriano, 2001). As a result of commercial interest various strains of Gracilaria spp. have been developed to produce higher agar yields (Bird, 1988). However, there is ‘No evidence’ to suggest that these strains of Gracilaria spp. have negatively affected natural populations.

Not relevant (NR)
NR
NR
NR
Help
Not relevant (NR)
NR
NR
NR
Help
No evidence (NEv)
NR
NR
NR
Help
Introduction or spread of invasive non-indigenous species [Show more]

Introduction or spread of invasive non-indigenous species

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

Evidence

Competition with invasive macroalgae may be a potential threat to this biotope (de Bettignies et al., 2021).  Potential invasives include Undaria pinnatifida and Sargassum muticum.  Sargassum muticum is a circumglobal invasive species (Engelen et al., 2015).  It is recorded (2015) from Norway to Morocco and into the Mediterranean in the eastern Atlantic and 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.  Although fertilization does not occur below 15 ppt and growth of germlings is limited below 10°C it can complete its life cycle as long as temperatures are over 8°C for at least four months of the year (Engelen et al., 2015).  However, 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.  But it has been recorded at 18 m or 30 m in the clear waters of California.  However, it is a poor competitor under low light and only develops dense canopies in shallow areas (Engelen et al., 2015).

Sargassum muticum was shown to replace and out-compete leathery, canopy-forming macroalgae such as Saccharina latissima, Halidrys siliquosa, and Fucus spp. and, to a lesser degree, understorey species such as Codium fragile, Chondrus crispus and Dictyota dichotoma in Limfjorden, Denmark between 1984 and 1997 (Staehr et al., 2000; Engelen et al., 2015; de Bettignies et al., 2021).  The invasion in Limfjorden had stabilized by 2005 although many of the native macroalgal species continued to decline (Engelen et al., 2015).  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 although wave exposure has been reported to restrict the growth and survival of Sargassum muticum (Staehr et al., 2000).  Viejo et al. (1995) reported that Sargassum muticum transplanted to wave exposed shores in Spain experienced >80% breakages within a month and that the growth of undamaged plants was significantly lower than that of plants on sheltered shores.  Similarly, Andrew & Viejo (1998) noted that Sargassum muticum was restricted to intertidal rockpools in wave exposed sites in the Bay of Biscay.

Strong & Dring (2011) used canopy removal experiments to investigate inter- and intra-species competition between Sargassum muticum and Saccharina latissima in the Dorn, Strangford Lough, N. Ireland.  The Dorn consists of tidal pools, very sheltered from wave action but with moderately strong tidal streams (1-2 knots).  Sargassum muticum grew better in mixed stands with Saccharina latissima than in the highest density monospecific stands examined.  However, the growth of Saccharina was not affected by the proportion of Sargassum in mixed stands.  They concluded that Saccharina was not impacted significantly by the alien species while Sargassum benefited from growth in mixed stands.  Experimental manipulation of subtidal algal canopies in San Juan Islands, Washington State, USA, showed that Sargassum muticum reduced the abundance of native macroalgae, including the kelp Laminaria bongardiana due to shadingHowever, experimental removal of Sargassum resulted in the recovery of native species within about one year (Britton-Simmons, 2004; Engelen et al., 2015).  The negative effects of Sargassum muticum on native macroalgae are mainly due to competition for light, rather than changes in nutrient availability, sedimentation or water flow (Britton-Simmons, 2004; Engelen et al., 2015).  

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).  Undaria pinnatifida 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 was first recorded in the UK in the Hamble Estuary in 1994 (Macleod et al., 2016).  It has since proliferated along UK coastlines.  One year after its discovery at the Queen Anne Battery marina, Plymouth, it had become a major fouling plant on pontoons (Minchin & Nunn, 2014).  Although initially restricted to artificial habitats, such as marinas and ports, it is now widespread in natural habitats in several areas, including Plymouth Sound.

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).  It is found predominantly in low intertidal to shallow subtidal habitats (Epstein et al., 2019b) and is significantly more abundant on artificial substrata compared to natural rocky substrata (Heiser et al., 2014; Epstein & Smale, 2018).  James (2017) suggested that Undaria pinnatifida could out-compete native species on artificial substrata (such as marinas and wharf structures).  In Plymouth, UK, De Leij et al. (2017) found that natural habitats with dense native macroalgal canopies, such as Laminaria hyperborea, Laminaria ochroleuca, Laminaria digitata, and Saccharina latissima had more resistance to Undaria pinnatifida invasion than disturbed or sparse canopies, due to limited space and light availability for Undaria pinnatifida recruits.  However, the dense canopies did not always prevent the invasion of Undaria pinnatifida as sporophytes were still recorded within dense Laminaria canopies, so canopy disturbance was not always required (De Leij et al., 2017; Epstein & Smale, 2018).

Undaria pinnatifida species behaves as a winter annual and recruitment occurs in winter followed by rapid growth through spring, maturity, and then senescence through summer, with only the microscopic life stages persisting through autumn.  It exhibits multiple dispersal strategies, such as short-range spore dispersal, and long-range dispersal as whole drift plants or fragments.  Undaria pinnatifida has spread rapidly across the UK and Europe, resulting in community-wide responses and impacts (Vaz-Pinto et al., 2014; Epstein & Smale, 2017).  Its impacts are complex and context-specific, depending on space, time, and taxa present in the introduced location (Epstein & Smale, 2017; Teagle et al., 2017; Tidbury, 2020).

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 can inhibit a broad range of habitats including – reefs; coastal brackish/saline lagoons; large shallow inlets and bays; estuaries; estuarine rocky habitats; natural or near natural estuary; coastal lagoons; and tidal rivers, estuaries, mudflats, sandflats and lagoons (James, 2017).  Undaria pinnatifida prefers sites sheltered with low wave exposure and weak tidal streams (Heiser et al., 2014; Epstein & Smale, 2018).  In natural habitats, Undaria pinnatifida was not recorded if the wave fetch was greater than 642 km but increased in abundance and cover in very sheltered sites (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 Saccharina latissima.  However, they reported that Undaria pinnatifida biomass was negatively related to Saccharina latissima in both intertidal and subtidal habitats. This was only statistically significant in subtidal habitats, which suggested that there was some competition between the two species (Epstein et al., 2019b). Heiser et al. (2014) surveyed 17 sites within Plymouth Sound, UK and found that Saccharina latissima was significantly more abundant at sites with Undaria pinnatifida with ca 5 Saccharina latissima individuals present per m², compared to ca 0.5 Saccharina latissima individuals per m² present at sites without Undaria pinnatifida.

Undaria pinnatifida has been reported to both co-exist with and out-compete Saccharina latissima (Farrell & Fletcher, 2006; Heiser et al., 2014; Epstein et al., 2019b). For example, in Torquay Marina, UK, Farrell & Fletcher (2006) completed a canopy removal experiment between 1996-2002. They reported that Saccharina latissima decreased in both control and treatment plots from ca 3 plants per 0.45 m² in 1996 to ca 1 plant per 0.45 m² in 1997 and had disappeared completely from pontoons by 2002.  This coincided with a significant increase in Undaria pinnatifida from zero plants per 0.45 m² in 1996 to ca 6 plants per 0.45 m² in 1997.  However, there was a slight decrease in Undaria pinnatifida in both control and treatment plots between 1997 and 1998.  By 2002, Undaria pinnatifida had recovered at control and treatment plots to ca 4-6 plants per 0.45 m² whereas Saccharina latissima had not.

Undaria pinnatifida was successfully eradicated on a sunken ship in Clatham 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.

The proliferation of Undaria pinnatifida and competition with native species may cause a reduction in local biodiversity (Valentine & Johnson, 2003; Vaz-Pinto et al., 2014; Arnold et al., 2016; Teagle, 2017; Tidbury, 2020).  A shift towards Undaria pinnatifida dominated beds could result in diminished epibiotic assemblages and lower local biodiversity compared with assemblages associated with native perennial kelp species, such as Laminaria spp. and Saccharina latissima (Arnold et al., 2016; Teagle et al., 2017).  In Plymouth, UK, Arnold et al. (2016) found that Undaria pinnatifida supported less than half the number of taxa and had no unique epibionts compared to Laminaria ochroleuca and Saccharina latissima (Arnold et al., 2016).

Sensitivity assessment.  The above evidence suggests that both Sargassum muticum and Undaria pinnatifida can both compete with and co-exist with Saccharina latissima, depending on local conditions.  For example, Undaria pinnatifida can out-compete Saccharina latissima in artificial habitats, such as in Torquay Marina, but within natural habitats, it can co-exist with native kelp species within its depth range (-1 to 4 m), as shown in Plymouth Sound, UK.  Similarly, Sargassum muticum out-competed Saccharina latissima in theLimfjorden but coexisted in the Dorn in Strangford Lough.

This Saccharina latissima dominated biotope (SS.SMp.KSwSS.SlatGraVS) is found at 0-10 m (JNCC, 2015), in full or variable salinity with moderately strong to negligible tidal streams and wave sheltered to extremely sheltered conditions.  The evidence above suggests that Undaria prefers sheltered conditions, with low tidal flow, in the shallow subtidal and sublittoral fringe (ca +1 to 4 m in depth), while Sargassum also prefers wave sheltered conditions and shallow water (ca 1 to 4 m depth).  Therefore, Undaria pinnatifida and Sargassum muticum are likely to threaten the most shallow (e.g. 0-5 m) examples of this biotope, where suitable hard substrata are available.  They may either co-exist with or out-compete Saccharina latissima, resulting in a potentially significant (25-75%) reduction in the abundance or extent of the native kelp and a possible decrease in the diversity of other macroalgae.  Therefore, resistance is assessed as ‘Low’ for most shallow, wave sheltered examples of the biotope, i.e. above 5 m in depth.  Both Sargassum and Undaria may colonize at 5-10 m but in low numbers so Saccharina will probably dominate.  Recovery after invasion by Sargassum or Undaria, although rapid, would require direct intervention (removal) so that resilience is assessed as ‘Very low’.  Hence, the sensitivity of shallow, sheltered, examples of the biotope is assessed as ‘High’.  Overall, confidence is assessed as ‘Low’ due to evidence of variation and site-specific nature of competition between native kelps, Sargassum muticum, and Undaria pinnatifida.

Low
Low
NR
NR
Help
Very Low
High
High
High
Help
High
Low
Low
Low
Help
Introduction of microbial pathogens [Show more]

Introduction of microbial pathogens

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

Evidence

Laminarians may be infected by the microscopic brown alga Streblonema aecidioides. Infected algae show symptoms of Streblonema disease, i.e. alterations of the blade and stipe ranging from dark spots to heavy deformations and completely crippled thalli Infection can reduce growth rates of host algae (Peters & Scaffelke, 1996). The marine fungi Eurychasma spp can also infect early life stages of Laminarians however the effects of infection are unknown (Müller et al., 1999).

Gracilaria gracilis is also susceptible to bacterial pathogens. Farmed and natural populations of Gracilaria gracilis within Saldanha Bay, South Africa have experienced a number of large die-offs since 1989, in which thalli have become bleached and/or rotten as a result of Pseudoalteromonas gracilis B9 infection (Schroeder et al., 2003).

Sensitivity assessment. Resistance to the pressure is considered ‘Low’ based on the potential susceptibility of Gracilaria gracilisand resilience is probably ‘High’. The sensitivity of this biotope to introduction of microbial pathogens is assessed as ‘Low’.

Low
Medium
Low
Medium
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High
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Removal of target species [Show more]

Removal of target species

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

Evidence

There has been recent commercial interest in Saccharina lattisma as a consumable called “sea vegetables” (Birket et al., 1998). However, Saccharina lattissima sporophytes are typically matured on ropes (Handå et al 2013) and not directly extracted from the seabed, as with Laminaria hyperborea (Christie et al., 1998). No evidence has been found for commercial extraction of Chorda filum. Gracilaria gracilis is commercially harvested for the production of agar (Marinho-Soriano, 2001). Over-exploitation of Gracilaria gracilis has, in some regions, resulted in depletion of natural stocks. As result, various culture techniques have been developed to increase yields. Natural stocks are still harvested however in some instances has been restricted to gathering beach cast (Martín et al., 2011). Furthermore removal of macro-algae canopy may also cause mortality of the understorey community (Hawkins & Harkin, 1985).

Sensitivity assessment. As a result of historical exploitation causing mass declines in Gracilaria gracilis resistance has been assessed as ‘None’, however confidence scores are marked as low. Resilience as ‘High’. Sensitivity has been assessed as ‘Medium’.

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Removal of non-target species [Show more]

Removal of non-target species

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

Evidence

Low level disturbances (e.g. solitary anchors) are unlikely to cause harm to the biotope as a whole, due to the impact’s small footprint. Thus evidence to assess the resistance of SS.SMp.KSwSS.SlatGraFS and SS.SMp.KSwSS.SlatGraVS to non-targeted removal is limited. It is assumed that incidental non-targeted catch (e.g. by trawls or dredges) could mobilise sediment, remove large kelp species, overturn boulders and cobbles and bury smaller seaweeds and cause high mortality within the affected area. Furthermore removal of macro-algae canopy may also cause mortality of the understorey community (Hawkins & Harkin, 1985).

Sensitivity assessment. Resistance has been assessed as ‘None’, Resilience as ‘High’. Sensitivity has been assessed as ‘Medium’.

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Citation

This review can be cited as:

Stamp, T.E., Williams, E., Lloyd, K.A., & Mardle, M.J., 2022. Saccharina latissima and Gracilaria gracilis with sponges and ascidians on variable salinity infralittoral sediment. In Tyler-Walters H. Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 29-03-2024]. Available from: https://marlin.ac.uk/habitat/detail/1056

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Last Updated: 23/06/2022