The Marine Life Information Network

Temperature increase (local)

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

Evidence

Schonbeck & Norton (1979) demonstrated that fucoids can increase tolerance in response to a gradual change in temperature through a process known as 'drought hardening'. However, acute changes in temperatures may cause damage to macroalgae and other species. Temperature ranges of species may not accurately describe their ability to withstand localized changes in temperature. However, they will display the limits of the species genetic ability to acclimatize to temperatures. The juvenile life stages of organisms can be less tolerant of environmental conditions than more mature stages.

Ascophyllum nodosum is found in the middle of its range in the British Isles, with populations in the North East Atlantic as far south as Portugal and extending north to the White Sea and Iceland and west into the Kattegat on the shores of Sweden. Ascophyllum nodosum is unlikely to be affected by a short-term change of 5°C, as it was not damaged during the unusually hot summer of 1983 when the average temperature was 8.3°C higher than normal (Hawkins & Hartnoll, 1985). Ascophyllum nodosum can tolerate certain levels of exposure as they are regularly exposed to rapid and short-term variations in temperature. Both exposure at low tide or rising tide on a sun-heated shore involves considerable temperature changes, and during winter the air temperature may be far below freezing point. The growth of Ascophyllum nodosum has been measured between 2.5 and 35°C with an optimum between 10 and 17°C (Strömgren, 1977). Ascophyllum nodosum can be damaged by thermal pollution if the water temperature remains above 24°C for several weeks (Lobban & Harrison, 1997), and temperatures exceeding 27-28°C cause direct mortality (Keser et al., 2005). Water temperature is an excellent predictor of gamete release in Ascophyllum (Bacon & Vadas, 1991). Consequently, changes in temperatures could impact on gamete release. Investigations into the tolerance of Ascophyllum nodosum germlings from Norway, to temperatures between 7°C -17°C, found that there was no difference in survival rates within the given range (Steen & Rueness, 2004). Germination of Ascophyllum nodosum has been recorded between the temperatures of 4°C and 23°C.

In the North East Atlantic, Fucus vesiculosus occurs from Northern Russia to Morocco (Powell, 1963). Fucus vesiculosus can tolerate temperatures as high as 30°C (Lüning, 1990) and did not show any sign of damage during the extremely hot UK summer of 1983 when average temperatures were 8°C hotter than normal (Hawkins & Hartnoll, 1985). Fucus vesiculosus also tolerates extended periods of freezing in the northern part of its range. Fucus serratus is found along the Atlantic coast of Europe from Svalbard to Portugal and on the shores of north-east America and is within its thermal range in the British Isles. Nielsen et al. (2014) found no negative effects on growth rates of adult Fucus serratus to water temperatures of 22°C (based on a laboratory experiment with specimens collected from Firth of Forth, Scotland) and Arrontes (1993) observed that Fucus serratus survived in laboratory experiments for 1 week at 25°C. Nielsen et al. (2014) did, however, report that germlings were negatively affected by increased temperature indicating that early life stages are more vulnerable than mature algae to this pressure.

Other species found within this biotope are probably tolerant of temperature changes at the benchmark level as they are widely distributed in the UK.  The balance of interactions between fucoids and barnacles changes with geographical location.  Warmer conditions further south than the British Isles favour greater penetration of barnacles into sheltered locations (Ballantine, 1961; cited in Raffaelli & Hawkins, 1996).  Warmer conditions are also likely to favour Chthamalus spp. rather than Semibalanus balanoides although a change of species will not alter the function of the biotope. Those species which are mobile, such as Carcinus maenas have the opportunity to move away from areas if physical conditions become too harsh.

Sensitivity assessment. The characterizing species Ascophyllum nodosum is found in the middle of its habitat range in the British Isles. Although the range of this species can extend down to Portugal, a short term acute temperature increase, leaving no time for acclimation, might be expected to result in some damage to or mortality of Ascophyllum, especially if the increase occurred during the summer months. However, the observations of Hawkins & Hartnoll (1985) suggest otherwise. Therefore, resistance is assessed as 'High' at the benchmark (an increase of 5°C for one month) in UK waters. Resilience is assessed as 'High' so that sensitivity is assessed as a 'Not sensitive' at the benchmark level.

Temperature decrease (local)

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

Evidence

Schonbeck & Norton (1979) demonstrated that fucoids can increase tolerance in response to a gradual change in temperature through a process known as 'drought hardening'. However, acute changes in temperatures may cause damage to macroalgae and other species. Temperature ranges of species may not accurately describe their ability to withstand localized changes in temperature. However, they will display the limits of the species genetic ability to acclimatize to temperatures. The juvenile life stages of organisms can be less tolerant of environmental conditions than more mature stages.

Ascophyllum nodosum is found in the middle of its range in the British Isles, with populations in the North East Atlantic as far south as Portugal and extending north to the White Sea and Iceland and west into the Kattegat on the shores of Sweden. The growth of Ascophyllum nodosum has been measured between 2.5 and 35°C with an optimum between 10 and 17°C (Strömgren, 1977). Water temperature is an excellent predictor of gamete release in Ascophyllum (Bacon & Vadas, 1991). Consequently, changes in temperatures could impact on gamete release. Investigations into the tolerance of Ascophyllum nodosum germlings from Norway, to temperatures between 7°C -17°C found that there was no difference in survival rates within the given range (Steen & Rueness, 2004). Germination of Ascophyllum nodosum has been recorded between the temperatures of 4°C and 23°C.  Ascophyllum nodosum was reported to survive freezing to -20°C (MacDonald et al., 1974, cited in Åberg, 1992a). 

In the North East Atlantic, Fucus vesiculosus occurs from Northern Russia to Morocco (Powell, 1963). Fucus vesiculosus can tolerate temperatures as high as 30°C (Lüning, 1990) and did not show any sign of damage during the extremely hot UK summer of 1983 when average temperatures were 8°C hotter than normal (Hawkins & Hartnoll, 1985). Fucus vesiculosus also tolerates extended periods of freezing in the northern part of its range. Fucus serratus is found along the Atlantic coast of Europe from Svalbard to Portugal and on the shores of north-east America and is within its thermal range in the British Isles. Lüning (1984) reported that Fucus serratus survived in the laboratory for a week a range temperature between 0°C and 25°C and placed this species in his 'cold temperature North Atlantic group'.

A large number of the species found within this biotope are found throughout the British Isles and are not on the edge of their range. Therefore, it is unlikely that a decrease in temperature is going to cause significant mortalities. In addition, mobile species such as Carcinus maenas have the opportunity to move away from areas if physical conditions become too harsh. Hence, these species may decrease in abundance.

Sensitivity assessment. The characterizing species Ascophyllum nodosum is found in the middle of its habitat range in the British Isles. It is unlikely to be affected by a short-term change of 5°C for one month or 2°C for a year in UK waters as it survives harsher winter conditions in northern waters of Iceland and in Sweden where populations are subject to ice scour (Åberg, 1992a,b; Ingólfsson & Hawkins, 2008). Therefore, resistance is assessed as 'High' at the benchmark (an increase of 5°C for one month) in UK waters. Resilience is assessed as 'High' so that sensitivity is assessed as a 'Not sensitive' at the benchmark level.

Salinity increase (local)

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

Evidence

Variation in salinity within saline lagoons is a characteristic of the habitat. In larger lagoons is it also possible to get different salinities within different areas of the lagoon itself. Therefore, all species found within saline lagoon communities must be able to tolerate salinity fluctuations. All lagoons are affected by short, medium and long-term changes in salinity (Bamber et al., 2001). Semi-diurnal tides cause differences in salinity over short periods, response to changes in rainfall can cause medium-term changes and seasonal rainfall and drought can cause longer-term salinity trends (Bamber et al., 2001). Intertidal macroalgae often experience large but short-term changes in salinities (Lobban & Harrison, 1997). Salinities within these habitats vary due to weather conditions such as rainfall at low tide and evaporation from rock pools causing hypersaline conditions on hot days. Intertidal shores within estuarine environments can also experience considerable short-term changes in salinities. However intertidal macroalgae tolerances to longer-term changes in salinities can be minimal and can quickly reduce photosynthetic abilities and cause mortality.

This biotope is recorded from 'variable' salinity (18 – 35 ppt) (Connor et al., 2004). Hence, an increase in salinity could make the conditions hypersaline. Little empirical evidence was found to assess how an increase in salinity at this benchmark would affect Ascophyllum nodosum. Baardseth, 1970 noted that Ascophyllum nodosum is euryhaline with a salinity tolerance of about 15 to 37 psu. Chock & Mathieson (1979) found Ascophyllum nodosum plants in the laboratory photosynthesised at salinities from 0 to 40 psu, although the long-term effects within this range were not evaluated. No information could be found on the effects of an increase in salinity on the reproductive cycle of Ascophyllum nodosum. 

A number of the species associated with this biotope can also be found within rockpools where hypersaline conditions can be found for short periods (Newell, 1979). Fucus vesiculosus is well adapted to cope with varying salinities and can grow in full saline to brackish conditions. Bäck et al. (1991) compared Fucus vesiculosus individuals from Atlantic and Baltic populations. Both populations were able to withstand a wide range of salinities in laboratory cultures. The Atlantic population showed better growth in higher salinities and virtually no growth at 5 ppt. Those individuals kept at 5 ppt mortality occurred after seven weeks. In contrast, the Baltic wracks grew better in conditions with lower salinities. But growth was negligible at the highest tested salinity of 45 ppt (Bäck et al., 1991). Growth rates for Fucus serratus are maximal at a salinity of 20 psu with the critical limit for recruitment set at 7 psu (Malm et al., 2001). Semibalanus balanoides can tolerate salinities between 12 and 50 psu; below and above this cirral activity ceases (Foster, 1970). Carcinus maenas is mobile can move to suitable conditions on the shore.

Sensitivity assessment. Although many species within this biotope would be able to cope with a short-term increase in salinity, long-term hypersaline conditions could cause mass mortalities of the biological community within this biotope.  However, no evidence on the effects of hypersaline conditions on Ascophyllum nodosum or its associated community was found. 

Salinity decrease (local)

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

Evidence

Variation in salinity within saline lagoons is a characteristic of the habitat. In larger lagoons is it also possible to get different salinities within different areas of the lagoon itself. Therefore, all species found within saline lagoon communities must be able to tolerate salinity fluctuations. All lagoons are affected by short, medium and long-term changes in salinity (Bamber et al., 2001). Semi-diurnal tides cause differences in salinity over short periods, response to changes in rainfall can cause medium-term changes and seasonal rainfall and drought can cause longer-term salinity trends (Bamber et al., 2001). Intertidal macroalgae often experience large but short-term changes in salinities (Lobban & Harrison, 1994). Salinities within these habitats vary due to weather conditions such as rainfall at low tide and evaporation from rock pools causing hypersaline conditions on hot days. Intertidal shores within estuarine environments can also experience considerable short-term changes in salinities. However intertidal macroalgae tolerances to longer-term changes in salinities are minimal and can quickly reduce photosynthetic abilities and cause mortality.

Ascophyllum nodosum is euryhaline with a salinity tolerance of about 15 to 37 psu (Baardseth, 1970). The species can also withstand periodic emersion in freshwater (Baardseth, 1970) and frequently inhabits estuaries where salinity is variable. Doty & Newhouse (1954) reported Ascophyllum nodosum from estuarine waters with a maximum salinity of 17.3 psu and a minimum of 0 psu. Chock & Mathieson (1979) found Ascophyllum nodosum plants in the laboratory photosynthesised at salinities from 0 to 40 psu, although the long-term effects within this range were not evaluated. In the Teign Estuary in South Devon, Ascophyllum nodosum inhabits areas subject to salinities as low as 8 psu (Laffoley & Hiscock, 1993). Investigations into the salinity tolerance of Ascophyllum nodosum in laboratory controlled conditions found that the photosynthetic capabilities of this species decreased with reduced salinities. Ascophyllum nodosum tolerated seven days at salinities of 5, and all samples died after 15 days at salinities of 5 (Connan & Stengel, 2011). There is some evidence to suggest that reduced salinities can influence the rate of receptacle maturation in fucoids (Munda, 1964). The rate of fructification in Ascophyllum nodosum has been measured to increase in diluted seawater (Munda, 1964).

Fucus vesiculosus is well adapted to cope with varying salinities and can grow in full saline to brackish conditions.  Fucus vesiculosus is the dominant large perennial seaweed in the Baltic Sea growing in salinities down to 4 psu (Kautsky, 1992). Bäck et al. (1991) compared Fucus vesiculosus individuals from Atlantic and the Baltic populations. Both populations were able to withstand a wide range of salinities in laboratory cultures, yet some differences were recorded. The Atlantic population showed better growth in higher salinities and virtually no growth at 5 ppt. Those individuals kept at 5 ppt mortality occurred after seven weeks. In contrast, the Baltic wracks grew better in conditions with lower salinities (Back et al., 1991).  Serrao et al. (1996a) found that lower salinities can negatively affect both the fertilization rates and recruitment success of Fucus vesiculosus. Serrao et al. (1996a) also concluded that the osmotic tolerances of Fucus vesiculosus gametes limit the species distribution in the Baltic Sea.

Growth rates for Fucus serratus are maximal at a salinity of 20 psu with the critical limit for recruitment set at 7 psu (Malm et al., 2001).  Sufficient salinity is essential for successful fertilization and germination in Fucus (e.g., Brawley, 1992; Serrão et al., 1999). Malm et al. (2001) found that fertilization success in Fucus serratus decreased substantially with strongly reduced salinity. Indeed the study found that fertilization success was 87% at 9 psu but declined to 5% at 6 psu (Malm et al., 2001). Reduced salinity also affects dispersal by decreasing the swimming performance of fucoid sperm (Serrão et al., 1996a). These studies show that low salinities limit the recruitment and fertilization success of fucoids. 

A number of the other species within the biotope can also be found within rockpools where hyposaline conditions can be found for short periods (Newell, 1979). Consequently, a decrease in salinity within the benchmark of this pressure may not cause significant mortalities. For example, Semibalanus balanoides can tolerate salinities between 12 and 50 psu, below and above this cirral activity ceases (Foster, 1970). Carcinus maenas is a mobile species and can move to suitable conditions on the shore.

Sensitivity assessment. This biotope is recorded from both variable (18 – 40 ppt) (Connor et al., 2004). A decrease in salinity at the benchmark would create a reduced salinity regime (18-30) for a year. As Ascophyllum nodosum occurs in estuarine conditions, inhabits areas subject to salinities as low as 8 psu, and can tolerate seven days at a salinity of 5 (Laffoley & Hiscock, 1993; Connan & Stengel, 2011) it is unlikely to suffer a reduction in abundance due to a reduction in salinity at the benchmark level.  Similarly, both of the characteristic Fucus spp. can tolerate low salinity. Therefore, resistance is assessed as 'High' at the benchmark so that resilience is assessed as 'High' and sensitivity as 'Not sensitive' at the benchmark level.  

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

Water motion is a key determinant of marine macroalgal ecology, influencing physiological rates and community structure (Hurd, 2000). Higher water flow rates increase mechanical stress on macroalgae by increasing drag. Fucoids are highly flexible but not physically robust and an increase in water flow could cause mechanical damage, breaking fronds or even dislodging whole algae from the substratum. Fucoids are, however, highly flexible and able to re-orientate their position in the water column to become more streamlined. This ability allows fucoids to reduce the relative velocity between algae and the surrounding water, thereby reducing drag and lift (Denny et al., 1998). Fucoids are permanently attached to the substratum and would not be able to re-attach if removed. Organisms living on the fronds and holdfasts will be washed away with the algae whereas free-living community components could find new habitat in surrounding areas. Wave exposure has been shown to limit the size of fucoids (Blanchette, 1997) as smaller individuals create less resistance to water movement, water flow likely exerts a very similar pressure on fucoids.

Fucus vesiculosus individuals of 10 cm or larger have been recorded to be completely removed at 7-8 m/s (Jonsson et al., 2006). Flow rates at which adult Ascophyllum nodosum are removed are not known. However, Thompson & Wernberg (2005) provide strong evidence of an increase in the break force required to remove algae with an increase in thallus size. Consequently, the force required to remove Ascophyllum nodosum from the shore is likely to be comparable to that of Fucus vesiculosus as both are large macroalgae with similar thallus sizes.

Propagule dispersal, fertilization, settlement, and recruitment are also influenced by water movement (Pearson & Brawley, 1996). An increase in water flow could have negative impacts on the reproductive success of Ascophyllum nodosum. Experiments on the effect of wave action on Ascophyllum nodosum showed that a low-velocity wave can remove 99% of 15-minute old zygotes from experimental tiles (Vadas et al., 1990). Further investigation with the use of refuges found that 75-90% of zygotes as old as four hours could be removed by a single wave. The attachment success of Ascophyllum nodosum was poor at current speeds of over 20 cm/s (Vadas et al., 1992). These studies show the need for periods of calm conditions for successful recruitment for Ascophyllum nodosum. An increase in the mean water flow could reduce the time during which attachment is possible. In addition, greater water flow can increase scour through increased sediment movement. Small life stages of macroalgae are likely to be affected by removing new recruits from the substratum and hence reducing successful recruitment (Devinny & Volse, 1978) (see ‘siltation’ pressures). Changes in water motion can thus strongly influence local distribution patterns of Fucus spp. (Ladah et al., 2008).

Sensitivity assessment. This lagoonal biotope (IR.LIR.Lag.AscSpAs) is recorded from weak (<1 kn, <0.5 m/s) to negligible tidal flow. A further decrease in water flow is unlikely to be relevant. Also, Ascophyllum nodosum is recorded in tide-swept conditions (e.g. LR.HLR.FT.AscT) in strong (1.5-3 m/s) to very strong (>3 m/s) water flow.  Therefore, a change in the current flow of 0.1-0.2 m/s is unlikely to have an impact on many examples of this biotope. Hence, resistance and resilience have been assessed as ‘High’ and the biotope is assessed as ‘Not Sensitive’ at the benchmark level.

Emergence regime changes

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

Evidence

Within the British Isles populations of Ascophyllum nodosum can suffer from bleaching and consequent mortality during exceptionally hot weather (Schonbeck & Norton, 1978, Hawkins & Hartnoll, 1985, Norton, 1985). However, these mortality events do not occur every year and tend to occur when the effects of unusually hot conditions combine with large tides and result in rapid changes that do not allow for macroalgae to acclimate (Raffaelli & Hawkins, 1996).

Stengel & Dring (1997) reported that growth rates in Ascophyllum nodosum decreased with height on the shore, correlating with an increase in environmental severity. Ascophyllum nodosum productivity is affected by desiccation when water loss exceeds 50% (Brinkhuis et al., 1976). Higher temperatures can increase the rate of desiccation and consequently lead to a loss of productivity, and eventually mortality (Keser et al., 1981). When Stengel & Dring (1997) transplanted Ascophyllum nodosum from the lower shore to the upper shore, 80% of the transplants died within 3 months.  In contrast, 100% of the individuals from the upper shore transplanted to the lower shore survived, as did all of the controls. The plants that survived transplantation to the upper shore acclimated to the conditions on the upper shore, yet their survival was determined by thallus morphology a predetermined genetic attribute which may be fixed (Stengel & Dring, 1997). Choi & Norton (2005) also carried out transplantation experiments and found that the growth rates of Ascophyllum nodosum decreased dramatically from the lower shore to the upper shore.

The southern and northern range limits of several intertidal macroalgae fall within Portugal. Lima et al. (2007) mapped the readjustment of 129 macroalgal ranges in relation to the change in air and sea temperatures observed within the north-eastern Atlantic over the past 50 years. Significant differences in distributions of algae were found, yet there was a disparity in the level of change found in the ranges of those of warm and cold adapted species. The species that were at the northern limit of their range in Portugal showed a greater change in distribution than the cold adapted species. Roughly half of the cold adapted species, including Ascophyllum nodosum, showed no significant change in their distribution. Lima et al. (2007) suggested that the cold adapted species had greater tolerance to adverse conditions for longer periods than the warm adapted species. 

Information regarding the effect of changes in the level of exposure on Ascophyllum nodosum germlings was not available. Germlings would be protected from desiccation stresses due to air exposure because of the protection provided to them by the fucoid canopy. Increases in temperature will be one of the effects changes in exposure will have on germlings. For further information refer to temperature pressure. Dense aggregations of algae can reduce the effect of more severe physical conditions such as those experienced with greater levels of exposure. Clumping enables organisms to retain moisture and reduce heat stress (Scrosati & DeWreede, 1998, Stafford & Davies, 2005).

Fucus vesiculosus can tolerate desiccation until the water content is reduced to ~30%. If desiccation occurs beyond this level, irreversible damage occurs. Individuals at the top of the shore probably live at the upper limit of their physiological tolerance and therefore, are likely to be unable to tolerate increased desiccation and would be displaced by more physiologically tolerant species. Tolerance to this pressure is likely to vary on a geographical scale. Gylle et al. (2009) found that Fucus vesiculosus populations that occur naturally in fully saline conditions had a higher emersion stress tolerance compared to brackish populations. Early life history stages are more susceptible to this pressure compared to adults (Henry & Van Alstyne, 2004). Germlings are, however, protected from desiccation by the canopy of adults. Brawley & Jonhnson (1991) showed that germling survival under the adult canopy was close to 100% whereas survival on the adjacent bare rock was close to 0% during exposure to aerial conditions.  Fucus serratus is more susceptible to desiccation than other Fucus species that are located further up the shore and subjected more frequently to aerial exposure (Schonbeck & Norton, 1978). In experiments, (Schonbeck & Norton, 1978) Fucus serratus did not survive transplantation further up the shore, e.g. in the Fucus spiralis belt. The critical water content for Fucus serratus is estimated at 40% with water losses past this point causing irreversible damage. Beer et al. (2014) found that Fucus serratus could not regain any positive photosynthetic rates after rehydrating from 10% water content.

Sensitivity assessment. This lagoonal biotope IR.LIR.Lag.AscSpAs occurs in the shallow sublittoral. However, lagoons are isolated water bodies and may experience changes in water level due to changes in inflow from the sea or rivers, periods of drought or human interference (e.g. drainage) and the pressure is considered relevant. Desiccation and the associated osmotic stress, especially when combined with high temperatures can cause mortalities (Pearson et al., 2009). The sensitivity of the characterizing species to emersion pressure will depend on the health and demography of individual populations, with germlings being most vulnerable life stage to this pressure. Ascophyllum nodosum has a level of resistance to an increase in emersion.  An increase in the emergence time for a year is likely to change in the height of the biotope on the shore, and exposure the biotope to intertidal conditions. It is likely to transition into an intertidal equivalent (e.g. LR.LLR.F.Asc) over time. Conversely, a decrease in emergence may allow the biotope to increase its extent up the shore, although its lower limit may be lost. Nevertheless, the change in the biotope within a year (the benchmark) may be small and the dominant canopy may not be lost. Therefore, resistance is assessed as 'Medium'. Hence, resilience is probably also 'Medium' and sensitivity is assessed as 'Medium'. 

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

An increase in wave exposure generally leads to a decrease in macroalgae abundance and size (Lewis, 1961, Stephenson & Stephenson, 1972, Hawkins et al., 1992, Jonsson et al., 2006).  Fucoids are highly flexible but not physically robust and an increase in wave exposure can cause mechanical damage, breaking fronds or even dislodging whole algae from the substratum. Ascophyllum nodosum is permanently attached to the substratum and would not be able to re-attach if removed. Organisms living on the fronds and holdfasts will be washed away with the algae whereas free-living community components could find new habitat in surrounding areas. Wave exposure has been shown to limit the size of fucoids (Blanchette, 1997) as smaller individuals create less resistance to waves. As exposure to waves increases the fucoid population will become dominated by small juvenile algae and dwarf forms of macroalgae which are more resistant to strong wave action. An increase in wave action beyond the tolerance of these fucoid species leads to a further increase in the abundance of robust fucoids, such as Fucus spiralis f. nana and red seaweeds, such as Corallina officinalis (Connor et al., 2004).

Ascophyllum nodosum cannot resist very heavy wave action so exposure to wave action is an important factor controlling the distribution of the species, and therefore this biotope. This biotope is found in sheltered to extremely sheltered conditions. Propagule dispersal, fertilization, settlement, and recruitment are also influenced by water movement (Pearson & Brawley, 1996). An increase in water flow due to wave exposure could have negative impacts on the reproductive success of Ascophyllum nodosum. Experiments on the effect of wave action on Ascophyllum nodosum showed that a low-velocity wave can remove 99% of 15-minute old zygotes from experimental tiles Vadas et al. (1990). Further investigation with the use of refuges found that 75-90% of zygotes as old as four hours could be removed by a single wave. Current speeds over 20 cm s^-1 make attachment success of Ascophyllum nodosum very poor (Vadas et al., 1992). These studies show the need for periods of calm conditions for successful recruitment for Ascophyllum nodosum. An increase in the mean wave exposure will reduce the time during which attachment is possible. In addition, greater wave action can increase scour through increased sediment movement. Small life stages of macroalgae are likely to be affected by removing new recruits from the substratum and hence reducing successful recruitment (Devinny & Volse, 1978) (see ‘siltation’ pressures). The other characterizing species are found in a range of wave exposures and unlikely to be directly affected. However, loss of the fucoid cover would result in major changes to the associated community, especially attached epifauna and understorey algae.

Sensitivity assessment. As this is an extremely sheltered biotope a further decrease in wave exposure is unlikely. An increase in wave action is likely to adversely affect fucoid cover, especially of Ascophyllum nodosum.  The biotope would probably be lost if wave exposure increased from e.g. sheltered to moderately exposed.  It is difficult to qualify a 3-5% change in significant wave height in terms of wave exposure, but the biotope is likely to have at least a ‘Medium’ resistance to an increase in wave exposure. Therefore, as resilience is probably ‘Medium’, sensitivity is also assessed as ‘Medium’. 

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.

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.

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.

Radionuclide contamination

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

Evidence

No evidence.

Introduction of other substances

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

Evidence

This pressure is Not assessed.

De-oxygenation

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

Evidence

Reduced oxygen concentrations have been shown to inhibit both photosynthesis and respiration in macroalgae (Kinne, 1977).  Despite this, macroalgae are thought to buffer the environmental conditions of low oxygen, thereby acting as a refuge for organisms in oxygen-depleted regions especially if the oxygen depletion is short-term (Frieder et al., 2012).  Reduced oxygen levels are likely to inhibit respiration whilst immersed, but it is unlikely to cause a loss of the macroalgae population directly.  As long as certain physical conditions are not exceeded, respiration and photosynthesis will be able to continue.

Although the macroalgae species within this biotope may not be negatively affected some of the associated fauna may be lost, causing a reduction in species richness.  Josefson & Widbom (1988) investigated the response of benthic macro and meiofauna to reduced dissolved oxygen levels in the bottom waters of a fjord. At dissolved oxygen concentrations of 0.21 mg/l, the macrofaunal community was eradicated and was not fully re-established 18 months after the hypoxic event. Meiofauna seemed, however, unaffected by deoxygenation.  Mobile species will be able to relocate to more optimal conditions, whereas immobile species such as barnacles are likely to be put under more stress by de-oxygenation.  Complete smothering caused by the Torrey Canyon oil spill appeared to have little impact on barnacle species; a few Semibalanus balanoides died, yet Chthamalus montagui seemed unaffected (Smith, 1968).  Semibalanus balanoides can respire anaerobically, so they can tolerate some reduction in oxygen concentration (Newell, 1979).  When placed in wet nitrogen, where oxygen stress is maximal and desiccation stress is low, Semibalanus balanoides have a mean survival time of 5 days (Barnes et al., 1963).

Sensitivity assessment.  The extremely wave sheltered conditions and the weak to very weak water flows which are characteristic of this biotope mean that water mixing is not very strong. Water retention rates within lagoons are also known to be high, with very low flushing times. Therefore, water movement within this area will not reverse any oxygen depletion quickly, possibly exacerbating any negative effects. The characterizing fucoid species may not be negatively affected by a decrease in oxygen within the water column at the benchmark level of this pressure.  However, some of the associated faunal community within this biotope may be negatively affected.  Mobile species such as the crab Carcinus maenas would relocate to conditions that were less physiologically taxing and would be able to return when the pressure abated.  Mysids may also be able to relocate. Those immobile species such as the barnacle Semibalanus balanoides may experience some mortality.  However, barnacles can completely recolonize within three years (Bennell, 1981). Therefore, as the fucoid canopy may survive a reduction in oxygen level to 2 mg/l or below for a week, resistance is assessed as ‘High’. Hence, resilience is assessed as ‘High’, and the biotope is assessed as 'Not sensitive'.

Nutrient enrichment

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

Evidence

woThe nutrient enrichment of a marine environment leads to organisms no longer being limited by the availability of certain nutrients. The consequent changes in ecosystem functions can lead to the progression of eutrophic symptoms (Bricker et al., 2008), changes in species diversity and evenness (Johnston & Roberts, 2009) decreases in dissolved oxygen and uncharacteristic microalgal blooms (Bricker et al., 1999, 2008).

Johnston & Roberts (2009) undertook a review and meta-analysis of the effect of contaminants on species richness and evenness in the marine environment. Of the 47 papers reviewed relating to nutrients as a contaminant, over 75% found that it had a negative impact on species diversity, <5% found increased diversity, and the remaining papers finding no detectable effect. Not all of the 47 papers considered the impact of nutrients on intertidal rocky shores. Yet this finding is still relevant as the meta-analysis revealed that the effects of marine pollutants on species diversity were ‘remarkably consistent’ between habitats (Johnston & Roberts, 2009). It was found that any single pollutant reduced species richness by 30-50% within any of the marine habitats considered (Johnston & Roberts, 2009). Throughout their investigation, there were only a few examples where species richness was increased due to the anthropogenic introduction of a contaminant. These examples were almost entirely from the introduction of nutrients, either from aquaculture or sewage outfalls. However, research into the impacts of nutrient enrichment from these sources on intertidal rocky shores often led to shores lacking species diversity and the domination by algae with fast growth rates (Abou-Aisha et al., 1995, Archambault et al., 2001, Arévalo et al., 2007, Diez et al., 2003, Littler & Murray, 1975).

Nutrient enrichment alters the selective environment by favouring fast-growing, ephemeral species such as Ulva lactuca and Ulva intestinalis (Berger et al., 2004, Kraufvelin, 2007). Rohde et al. (2008) found that both free growing filamentous algae and epiphytic microalgae can increase in abundance with nutrient enrichment. This stimulation of annual ephemerals may accentuate the competition for light and space and hinder perennial species development or harm their recruitment (Berger et al., 2003; Kraufvelin et al., 2007). Nutrient enrichment can also enhance fouling of Fucus fronds by biofilms (Olsenz, 2011). Nutrient enriched environments can not only increase algae abundance but the abundance of grazing species (Kraufvelin, 2007).

White et al. (2011) investigated the effects of nutrient effluent from land-based finfish farms on the morphologies of Ascophyllum nodosum in the vicinity of the outfall pipes. It was estimated that the nitrogen effluent from the farm was 1500 kg/yr. The background levels of nitrite at the test site were 300 μM.  In comparison, the ambient nitrite levels in south-west Nova Scotia are 3 μM (White et al., 2011). Ascophyllum nodosum at the test sites were found to be younger than those at the control sites, but significantly larger. This experiment suggested that nutrient effluent could have positive impacts on Ascophyllum nodosum. Yet it must be noted that the effect of the effluent on the rest of the biological community was not studied. Major declines of Fucus vesiculosus were reported from all over the Baltic Sea associated with eutrophication from nutrient enrichment (Kautsky et al., 1986). In experimental rocky shore communities, Krauflin et al. (2006) found only minor effects on the fucoid community structure as a response to high nutrient levels during the first three years of the experiment. However, during the 4^th year of exposure Fucus serratus started to decline and population consequently crashed in the 5^th year. The study observed full recovery of the algal canopy and animal community in less than two years after conditions returned to normal (Krauflin et al., 2006). 

Changes in community composition on intertidal rocky shores can happen rapidly, and fast-growing ephemeral species can become established quickly in the presence of higher concentrations of nutrients. The establishment and growth of these species are not controlled by wave exposure (Kraufvelin, 2007). However, even though these fast-growing ephemeral species can become well established quickly, healthy communities on intertidal rocky shores can survive long periods, and maintain ecological function after these species have become established (Bokn et al., 2002, 2003; Karez et al.,2004; Kraufvelin et al., 2006; Kraufvelin, 2007).

Sensitivity assessment. A slight increase in nutrients may enhance growth rates but high nutrient concentrations could lead to the overgrowth of the algae by ephemeral green algae and an increase in the number of grazers. If the biotope is well established and in a healthy state the biotope could persist. However, the biotope is ‘Not sensitive’ at the pressure benchmark that assumes compliance with good status as defined by the WFD.

Organic enrichment

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

Evidence

The organic enrichment of a marine environment at this pressure benchmark leads to organisms no longer being limited by the availability of organic carbon. The consequent changes in ecosystem functions can lead to the progression of eutrophic symptoms (Bricker et al., 2008), changes in species diversity and evenness (Johnston & Roberts, 2009) and decreases in dissolved oxygen and uncharacteristic microalgae blooms (Bricker et al., 1999, 2008).

Johnston & Roberts (2009) undertook a review and meta-analysis of the effect of contaminants on species richness and evenness in the marine environment. Of the 49 papers reviewed relating to sewage as a contaminant, over 70% found that it had a negative impact on species diversity, <5% found increased diversity, and the remaining papers finding no detectable effect. Not all of the 49 papers considered the impact of sewage on intertidal rocky shores. Yet this finding is still relevant as the meta-analysis revealed that the effects of marine pollutants on species diversity were ‘remarkably consistent’ between habitats (Johnston & Roberts, 2009). It was found that any single pollutant reduced species richness by 30-50% within any of the marine habitats considered (Johnston & Roberts, 2009). Throughout their investigation, there were only a few examples where species richness was increased due to the anthropogenic introduction of a contaminant. These examples were almost entirely from the introduction of nutrients, either from aquaculture or sewage outfalls. However, research into the impacts of organic enrichment from these sources on intertidal rocky shores often lead to shores lacking species diversity and the domination by algae with fast growth rates (Abou-Aisha et al., 1995, Archambault et al., 2001, Arévalo et al., 2007, Diez et al., 2003, Littler & Murray, 1975).

Nutrient enrichment alters the selective environment by favouring fast-growing, ephemeral species such as Ulva lactuca and Ulva intestinalis (Berger et al., 2004, Kraufvelin, 2007). Rohde et al., (2008) found that both free growing filamentous algae and epiphytic microalgae can increase in abundance with nutrient enrichment. This stimulation of annual ephemerals may accentuate the competition for light and space and hinder perennial species development or harm their recruitment (Berger et al., 2003; Kraufvelin et al., 2007). Nutrient enrichment can also enhance fouling of fucoid fronds by biofilms (Olsenz, 2011). Nutrient enriched environments cannot only increase algae abundance but the abundance of grazing species (Kraufvelin, 2007). Bellgrove et al. (2010) found that coralline turfs out-competed fucoids at a site associated with organic enrichment caused by an ocean sewage outfall.

White et al. (2011) investigated the effects of nutrient effluent from land-based finfish farms on the morphologies of Ascophyllum nodosum in the vicinity of the outfall pipes. It was estimated that the nitrogen effluent from the farm was 1500 kg/yr. The background levels of nitrite at the test site were 300 μM.  In comparison, the ambient nitrite levels in south-west Nova Scotia are 3 μM (White et al., 2011). Ascophyllum nodosum at the test sites were found to be younger than those at the control sites, but significantly larger. This experiment suggested that nutrient effluent could have positive impacts on Ascophyllum nodosum. Yet it must be noted that the effect of the effluent on the rest of the biological community was not studied. Major declines of Fucus vesiculosus were reported from all over the Baltic Sea associated with eutrophication from nutrient enrichment (Kautsky et al., 1986). In experimental rocky shore communities, Krauflin et al. (2006) found only minor effects on the fucoid community structure as a response to high nutrient levels during the first three years of the experiment. However, during the 4^th year of exposure Fucus serratus started to decline and population consequently crashed in the 5^th year. The study observed full recovery of the algal canopy and animal community in less than two years after conditions returned to normal (Krauflin et al., 2006). 

Changes in community composition on intertidal rocky shores can happen rapidly, and fast-growing ephemeral species can become established quickly in the presence of higher concentrations of nutrients. The establishment and growth of these species are not controlled by wave exposure (Kraufvelin, 2007). However, even though these fast-growing ephemeral species can become well established quickly, healthy communities on intertidal rocky shores can survive long periods of time, and maintain ecological function after these species have become established (Bokn et al., 2002, 2003, Karez et al.,2004; Kraufvelin et al., 2006; Kraufvelin, 2007).

Sensitivity assessment. Little empirical evidence was found to support an assessment of this biotope at this benchmark. Due to the negative impacts that can be experienced with the introduction of excess organic carbon both resistance and resilience have been assessed as ‘Medium’. This gives an overall sensitivity assessment of ‘Medium’.

Physical loss (to land or freshwater habitat)

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

Evidence

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

Physical change (to another seabed type)

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

Evidence

This biotope occurs on rock substratum so that a change to sedimentary or soft rock substratum would lead to the direct loss of suitable attachment areas. This change in substratum would result in the loss of the characterizing species Ascophyllum nodosum along with other species found within the associated community of this biotope, and reclassification of the biotope. Therefore, resistance is assessed as ‘None’. As this pressure represents a permanent change, recovery is impossible as a suitable substratum for the biological community of this biotope is lacking. Hence, resilience is assessed as ‘Very low’ and sensitivity is assessed as ‘High’. Although no specific evidence is described confidence in this assessment is ‘High’, due to the incontrovertible nature of this pressure.  

Physical change (to another sediment type)

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

Evidence

Not relevant.

Habitat structure changes - removal of substratum (extraction)

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

Evidence

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

Abrasion / disturbance of the surface of the substratum or seabed

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

Evidence

No studies of the effects of trampling or netting on lagoons were found but studies of the effects on emergent algal communities are probably indicative.  Trampling on the rocky shore has been observed to reduce fucoid cover which decreased the microhabitat available for epiphytic species, increased bare space and increased the cover of opportunistic species such as Ulva (Fletcher & Frid, 1996). This biotope is found in the mid intertidal shore; an area easily accessible by humans, especially at low tide. Fucoids are intolerant of abrasion from human trampling, which has been reported to reduce the cover of seaweeds on a shore (Holt et al., 1997; Tyler-Walters & Arnold, 2008).

Brosnan (1993) investigated the effect of trampling on a number of algal species, including Fucus vesiculosus, on an intertidal rocky shore in Oregon. The effects of 250 tramples per plot, once a month for a year were recorded. Abundances of algae in each plot were reduced from 80% to 35% within a month of the introduction of the pressure and remained low for the remainder of the experiment.  As few as 20 steps / m² on stations on an intertidal rocky shore in northeast England were sufficient to reduce the abundance of fucoids (Fletcher & Frid, 1996). A trampling intensity of 20 steps per m² per spring tide could be exceeded by only five visitors taking the same route out and back again across the rocky shore in each spring tide. This reduction in the complexity of the algae community, in turn, reduced the microhabitat available for epiphytic species. Trampling pressure can thus result in an increase in the area of bare rock on the shore (Hill et al., 1998). Chronic trampling can affect community structure with shores becoming dominated by algal turf or crusts (Tyler-Walters & Arnold, 2008).

Pinn & Rodgers (2005) compared the biological communities found on two intertidal rocky shore ledges in Dorset. They found that the ledge that had a higher number of visitors had few branching algal species, including fucoids, but had greater abundances of crustose and ephemeral species (Pinn & Rodgers, 2005). The densities of fucoids were recorded from the intertidal rocky shore at Wembury, Devon in 1930 (Colman, 1933) and 1973 (Boalch et al., 1974). Boalch et al. (1974) found a reduction in fucoids on the shore at Wembury (accessed by the public) and that the average frond length of Ascophyllum nodosum was smaller.

Fucus vesiculosus can generate vegetative regrowth in response to wounding from physical disturbance. McCook & Chapman (1992) experimentally tested the recovery of damaged Fucus vesiculosus. The study found that vegetative sprouting of Fucus vesiculosus holdfasts made a significant addition to the regrowth of the canopy, even when holdfasts were cut to less than 2 mm tissue thickness. Four months after cutting, sprouts ranged from microscopic buds to shoots about 10 cm long with mature shoots widespread after 12 months. Vegetative regrowth in response to wounding has been suggested as an important means of recovery from population losses (McLachan & Chen, 1972).

Ascophyllum nodosum seems to be particularly intolerant of damage from trampling (Flavell, unpublished; cited in Holt et al., 1997), as its length means it is more likely that the thallus is ‘cut’ between a footstep and sharp rock (Boalch et al., 1974, Tyler-Walters & Arnold, 2008). Araujo et al. (2009) found that trampling negatively affected both Ascophyllum nodosum abundances and reduced understorey species and promoted the colonization by ephemeral green algae. However, within a year of the disturbance event, Fucus vesiculosus had become the dominant canopy-forming species, replacing a pre-disturbance Ascophyllum nodosum community. The replacement of Ascophyllum nodosum with Fucus vesiculosus may have been due to the poor recovery rate of Ascophyllum nodosum. The increase in abundance suggests the competitive superiority of Fucus vesiculosus individuals in occupying newly available space in the disturbed patches. Similar results were found by Jenkins et al. (2004), Cervin et al. (2005) and Araujo et al. (2012) with Fucus vesiculosus outcompeting Ascophyllum nodosum after small scale disturbances. Rita et al. (2012) also undertook experiments on the effect of trampling on Ascophyllum nodosum and its associated communities. They concluded that trampling caused significant damage to both the macroalgae and the understorey communities, which had not recovered within five years of the initial experiment.

Sensitivity assessment. Abrasion of the substratum will cause a reduction in the abundances of Ascophyllum nodosum, as well as other species found in the associated community. Therefore, the resistance is assessed as ‘Low’. Experiments undertaken on the trampling effects on Ascophyllum nodosum have shown that for the community to return to its pre-experimental state can take over 12 years.  Hence, resilience is assessed as ‘Low’ and sensitivity as ‘High’. 

Penetration or disturbance of the substratum subsurface

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

Evidence

The species characterizing this biotope group are epifauna or epiflora occurring on hard rock, which is resistant to subsurface penetration.  Therefore, ‘penetration’ is 'Not relevant'. The assessment for abrasion at the surface only is, therefore, considered to equally represent sensitivity to this pressure’. Please refer to ‘abrasion’ above.

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

Light is an essential resource for all photoautotrophic organisms and a change in turbidity would affect light availability to photosynthesising organisms during immersion which could result in reduced biomass of plants. Changes in the suspended sediment load can change the levels of scour and alter the abundances of certain species. Greater levels of suspended particulate matter may also increase the amount of material which is falling out of suspension, which could consequently smother organisms (see siltation pressures). An increase in turbidity would alter the light available for photosynthesis during immersion. The shallow water depth within this biotope means that although light attenuation will be greater, the change in turbidity at this pressure benchmark will still allow light to penetrate to the depth at which the algae are found. Ascophyllum nodosum will also be able to continue to photosynthesize at low tide when the plants are emersed, as long as the plant has sufficient water content (Beer & Kautsky, 1992).

Daly & Mathieson (1977) found that Ascophyllum nodosum was completely absent from an intertidal rocky shore which was subject to a high level of scour from sand movement. The lack of Ascophyllum nodosum from this shore was particularly conspicuous due to the high abundance of the species on a nearby rocky shore with very similar conditions, except for the level of suspended sediment. Ascophyllum nodosum is not likely to be directly intolerant of a decrease in suspended sediment because the species is a primary producer.  The distribution of both Ascophyllum nodosum and Fucus vesiculosus occur along a depth gradient that strongly correlates with light penetration. In areas with low sedimentation, Fucus vesiculosus can survive down to 9-10 m depth (Eriksson & Bergstrom, 2005). Changes in suspended solids affecting water clarity may have a direct impact on the photosynthesising capabilities of Fucus vesiculosus.  Kõuts et al. (2006) found decreases in light intensity in the vicinity of the dredging site resulted in the net decline of Fucus vesiculosus biomass. A decrease in light penetration in the Kiel Fjord caused by an increase in phytoplankton density and shading from filamentous algae caused an upward shift of the lower depth limit of Fucus vesiculosus (Rohde et al., 2008). In extreme turbidity, such as found in the Bristol Channel, Fucus serratus is excluded from the bottom of the intertidal (below 2 m above chart datum) due to the lack of light for sustained growth (Chapman, 1995). 

Scour caused by increased sediment in suspension can cause mortality to many of the other species found within this biotope. For example, Daly & Mathieson, (1977) found that Semibalanus balanoides could be totally removed from a shore if scour is severe enough. A reduction in light levels due to an increase in the level of suspended sediment will not have a negative impact on the fauna within this biotope, and it is unlikely to have a significant negative impact on the other flora species, due to the intertidal nature of the biotope. An increase in levels of suspended sediment could be beneficial to filter-feeding organisms.

Sensitivity assessment. This biotope is found in the shallow subtidal and is permanently immersed. An increase in suspended sediment may cause the biotope to be lost at its lowest extent due to a decrease in light levels. The levels of water movement within this biotope, through water flow and wave exposure, are very unlikely to cause any significant damage through scour. Hence, the resistance and resilience of this biotope have been assessed as ‘Medium’ so that the sensitivity to this pressure is assessed as ‘Medium’.

Smothering and siltation rate changes (light)

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

Evidence

A discrete event where sediment inundates this biotope to 5 cm will have very different effects on the characterizing species and the associated community depending on the state of the tide. High tide will mean that both of the characterizing species will be vertical in the water column, meaning only a small proportion of the stipe and holdfast will be smothered, leaving the fronds sediment free, and able to continue photosynthesising. In contrast, if the tide is out then fronds of the characterizing fucoid canopy will be flat on the substratum and will be smothered by the sediment deposit. The level of water flow caused by tidal movements and wave exposure within this biotope will mean that the sediment won’t be removed from the shore quickly. Smothering will prevent photosynthesis resulting in reduced growth and eventually death.

However, germlings are likely to be smothered and killed in both scenarios and are inherently most susceptible to this pressure. Indeed early life stages are smaller in size than adults and are thus most vulnerable to this pressure as even a small load of added sediment will lead to the complete burial. Sediment deposition can reduce macroalgal recruitment by (1) reducing the amount of substratum available for attachment of propagules; (2) scour, removing attached juveniles and (3) burial, altering the light and/or the chemical micro-environment (Devinny & Volse, 1978, Eriksson & Johansson, 2003). Eriksson & Johansson (2003) found that sedimentation had a significant negative effect on the recruitment success of Fucus vesiculosus. Even fine sediment fractions can reduce the successful attachment of fucoids by as much as 90% (Schiel & Foster, 2006).

Ascophyllum nodosum is intolerant of sediment movement. Daly & Mathieson (1977) compared two rocky shores that were similar except for the level of sediment movement experienced on the shore. The shore with more sediment movement was devoid of Ascophyllum nodosum. Smothering will cause direct mortalities in the associated community, notably of the filter-feeding sessile organisms unable to clear their feeding appendages or relocate. Airoldi & Hawkins (2007) found that Patella vulgata reduces its feeding activity by 35% with just 1 mm of sediment over the substratum (equivalent to 50 mg/cm²). At 200 mg/cm² mortality occurred. It is possible that 5 cm of sand may create similar mortality events to other grazing organisms, as not only will they be weighted down by sand but food availability will also be restricted.

Sensitivity assessment. Ascophyllum nodosum adults are sediment intolerant, and germlings of Ascophyllum nodosum are intolerant of even small levels of sediment. Many of the smaller species found within the associated community will be smothered by 5 cm.  The level of water movement within this biotope is small and, consequently, deposited sediment will persist over a number of tides before it is all entrained and removed.  This is likely to cause some damage to the characterizing species and the other associated species.  Therefore, resistance is assessed as ‘Medium’. Resilience is probably 'Low' so that sensitivity is assessed as ‘Medium’ at the level of the benchmark.

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

Several studies found that increasing the vertical sediment burden negatively impact fucoids survival and associated communities.  At the level of the benchmark (30 cm of fine material added to the seabed in a single event), smothering is likely to result in mortalities of understorey algae, invertebrate grazers and young (germling) fucoids. Water movement will remove sediment but within this biotope is it likely to take a number of tidal cycles.  Resistance and resilience are assessed as  ‘Low’ and sensitivity as ‘High’ to siltation at the pressure benchmark.

Litter

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

Evidence

Not assessed.

Electromagnetic changes

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

Evidence

No evidence.

Underwater noise changes

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

Evidence

Species characterizing this habitat do not have hearing perception but vibrations may cause an impact, however, no studies exist to support an assessment.

Introduction of light or shading

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

Evidence

Increased levels of diffuse irradiation correlate with increased growth in macroalgae (Aguilaria et al., 1999). Levels of diffuse irradiation increase in summer, and with a decrease in latitude. As Ascophyllum nodosum is found in the middle its natural range in the British Isles an increase in the level of diffuse irradiation will not negatively impact the species or the biotope. However, it is not clear how these findings may reflect changes in light levels from artificial sources, and whether observable changes would occur at the population level as a result.

Cervin et al. (2005) noted that loss of canopy and degradation of the underlying turf promoted the recruitment of Ascophyllum nodosum to experimental plots. They also reported that Ascophyllum recruits had low growth rates in shade, under the canopy, that prevented the development of mature Ascophyllum plants. The modal size of Ascophyllum plants without canopy after six years was over twice that of individuals that grew under an intact canopy and the maximum size was six times greater (Cervin et al., 2005). It is possible that artificial shading, e.g. from a jetty, could slow the growth of Ascophyllum and decrease its ability to compensate for grazing or its ability to out-compete other fucoids.  However, no evidence was found to support an assessment. 

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

This pressure is considered applicable to mobile species, e.g. fish and marine mammals rather than seabed habitats. Physical and hydrographic barriers may limit propagule dispersal.  But propagule dispersal is not considered under the pressure definition and benchmark. Therefore this pressure is considered ‘Not Relevant’ for this biotope.

Death or injury by collision

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

Evidence

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

Visual disturbance

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

Evidence

Not relevant.

Genetic modification & translocation of indigenous species

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

Evidence

Key characterizing species within this biotope are not cultivated or translocated. This pressure is, therefore, considered ‘Not relevant’ to this biotope.

Introduction or spread of invasive non-indigenous species

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

Evidence

Thompson & Schiel (2012) found that native fucoids show high resistance to invasions by the Japanese kelp Undaria pinnatifida. However, the cover of Fucus vesiculosus was inversely correlated with the cover of the invasive Sargassum muticum indicating competitive interaction between the two species (Stæhr et al., 2000). Stæhr et al. (2000) determined that the invasion of Sargassum muticum could affect local algal communities through competition mainly for light and space.

Gracilaria vermiculophylla is suggested to be one of the most successful marine non-native species (Kim et al., 2010; Sfriso et al., 2010 cited in Thomsen et al., 2013). This species invades wave sheltered, shallow water areas, and have been found in biotopes naturally dominated by fucoid canopies (Weinberger et al., 2008). To date, Gracilaria vermiculophylla has only been recorded in Northern Ireland, and not on mainland Britain. The introduction of this species to intertidal rocky shores around the British Isles could have negative impacts on native fucoid biotopes and could become relevant to this specific biotope.

Sensitivity assessment. Fucoid species have been negatively affected by both the direct and indirect consequences of INNS being present.  However, no evidence was found on the impacts of INNS on Ascophyllum nodosum within this biotope.  Literature for this pressure should be revisited.

Introduction of microbial pathogens

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

Evidence

No evidence.

Removal of target species

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

Evidence

Seaweeds have been collected from the middle of the 16th century for the iodine industry. Modern industrial uses for seaweed are extensive and include fertilizer, animal feed, alginate extracts (Phillippi et al., 2014), water treatment, and human food and health supplements (Bixler & Porse, 2010). The characteristic fucoid algae within this biotope are commercially collected. These commercial harvests remove seaweed canopies which have important effects on the wider ecosystem. Due to the intolerance of macroalgae communities to human exploitation, the European Union put in place a framework to regulate the exploitation of algae establishing an organic label that implies that ‘harvest shall not cause any impact on ecosystems’ (no. 710/2009 and 834/2007).

Stagnol et al. (2013) investigated the effects of commercial harvesting of intertidal fucoids on ecosystem biodiversity and functioning. The study found that the removal of the macroalgae canopy affected the metabolic flux of the area. Flows from primary production and community respiration were lower on the impacted area as the removal of the canopy caused changes in temperature and humidity conditions (Stagnol et al., 2013). Bertness et al. (1999) found that the presence of an Ascophyllum nodosum canopy reduced maximum daily rock temperatures by 5-10°C. It was also reported that water loss via evaporation was an order of magnitude less than that in areas where the fucoid canopy had been removed (Bertness et al., 1999). Stagnol et al. (2013) found that suspension feeders were the most affected by the canopy removal as canopy-forming algae are crucial habitats for these species. Other studies confirm that loss of canopy had both short and long-term consequences for benthic community diversity resulting in shifts in community composition and a loss of ecosystem functioning such as primary productivity (Lilley & Schiel, 2006; Gollety et al., 2008).

Studies on the effects of commercial harvesting on the faunal communities associated with Ascophyllum nodosum have found that removing this key species can reduce abundances of epifauna found on the un-harvested biomass (Jarvis & Seed, 1996, Johnson & Scheibling, 1987; taken from Phillippi et al., 2014). Changes Ascophyllum nodosum have also been found to affect the large, mobile fauna such as crabs or grazing gastropods (Bertness et al., 1999; Fegley, 2001; Jenkins et al., 1999, 2004, Phillipi et al., 2014; Pocklington et al., 2018). Phillippi et al. (2014) replicated commercial harvesting techniques in Maine, USA where Ascophyllum nodosum fronds were removed 40.6 cm from the holdfast and the lowest lateral branch must remain with the holdfast (DMR, 2009). The experiment looked specifically at the effect of canopy reduction on infaunal species living within the soft sediments within intertidal rocky shores where Ascophyllum nodosum was present. The experiment found that invertebrate species found living on and within sediments were not negatively affected by the harvesting activity (Phillippi et al., 2014). However, Pocklington et al. (2018) examined community disturbance after removal of 100%, 50%, 245% and 0% of Ascophyllum nodosum fronds (but not holdfasts). They concluded that a pulse disturbance (frond removal) of 50% loss of fronds increased the temperature under the canopy significantly and decreased the abundance of mobile invertebrates such as Littorina obtusata. Sessile taxa such as Osmundia pinnatifida and encrusting corallines could withstand a 75% loss of fronds but declined by half if 100% were removed.  

Keser et al. (1981) recorded the levels of re-growth exhibited by Ascophyllum nodosum and Fucus vesiculosus after experimental harvesting in Maine. Harvesting was simulated by cutting fronds to three different lengths, that is, frond removed to the holdfast, to 15 cm from the holdfast and to 25 cm from the holdfast. Subsequent harvesting was repeated annually for three years. The experiment was carried out at eight sites, six of which were in sheltered areas. Re-growth of Ascophyllum nodosum was found to be dependent on; the age structure of the population; the extent and pattern of branching with a clump; the presence or absence of grazers (importantly Littorina littorea), and the environmental conditions. Recovery was found to be more rapid in estuaries (Keser et al., 1981). Of the fronds which that were cut back to the holdfast, only those within sheltered, estuarine and grazer free conditions showed any re-growth. More mature Ascophyllum nodosum fronds cut back to 15 cm and 25 cm within a sheltered site showed some re-growth, however, there were high rates of mortality.  The lack of re-growth was suggested to be caused by a lack of functional growing points found towards the bottom of the frond in older individuals. Almost all (95%) of young Ascophyllum nodosum individuals cut back to 15 cm and 25 cm regrew. In almost all populations measured within the experiment, repeat harvests resulted in lower biomass yields (Keser et al., 1981). Printz (1959) also carried out harvesting experiments where fronds were cut back to 25 cm, 15 cm and 5 cm from the holdfast. Individuals that had been cut back to 25 cm had an ‘abundance of new shoots’ and had grown to 30-35 cm in length after a year.  Individuals that had been trimmed back to 5 cm showed almost no change a year after the harvesting event. When the 5 cm individuals where re-visited three years after the harvesting event they were still almost unaltered. The reasons for the lack of re-growth were attributed to the lack of regenerative tissue found in the older flesh further down the thallus (Printz, 1959). There is considerable evidence to suggest that if Ascophyllum nodosum fronds are cut higher up the thallus recovery times are reduced considerably to two to three years (Ang et al., 1996; Fegley, 2001; Keser et al., 1981; Sharp, 1987; Ugarte et al., 2006; cited in Phillippi et al., 2014).  However, Keser et al. (1981) noted that repeated annual harvest reduced biomass and suggested staggering annual harvest between sites to prevent large-scale destruction of the resource. 

Sensitivity assessment. The removal of Ascophyllum nodosum canopy will significantly change the community composition of the biotope. The quantity of biomass removed from the shore and the regularity of removal will all affect how quickly the biotope will be able to recover. Ascophyllum nodosum probably has a ‘Low’ resistance to removal as it is easy to locate and has no escape strategy. However, resilience to harvesting (the removal of fronds above the holdfast) is probably ‘Medium’ (2-10 years) depending on the cut length, site, grazing pressure and age-size composition of the population (Keser et al., 1981; Phillippi et al., 2014). Therefore, sensitivity is assessed as ‘Medium’.

Removal of non-target species

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

Evidence

Direct, physical impacts from harvesting, trampling or removal are assessed through the abrasion and penetration of the seabed pressures.  This pressure focuses on the biological effects of the incidental (accidental) removal of a proportion of the dominant members of the community. Loss of the Ascophyllum canopy has been shown to alter the understorey community of sessile and mobile inveretebrates and red algal turf species significantly (Jenkins et al., 1999, 2004; Cervin et al., 2005; Phillippi et al., 2014; Pocklington et al., 2018). Subsequent recovery of the community results in intermediary communities, dominated by space and grazers or dominated by other fucoids, that may not correspond to this biotope.

Sensitivity assessment. Removal of a large percentage of the dominant characterizing species would alter the character of the biotope. The resistance to incidental removal is assessed as ‘Low’ due to the easy accessibility of the biotopes location and the inability of the species to evade removal. Therefore, resilience is assessed as ‘Low’ and sensitivity is as ‘High’.