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.

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). 

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

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 fully saline conditions (30 -40 ppt) (Connor et al., 2004). Consequently, 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). Consequently, an increase in salinity within the benchmark may not cause negative impacts. 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

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).

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) and fully saline conditions (30 -40 ppt) (Connor et al., 2004). A decrease in salinity at the benchmark would create a reduced salinity regime (18-30) for a period of 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.  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 biotope (LR.LLR.F.Asc) is recorded tidal currents ranging from 1 - 3 knots (0.5 – 1.5 m/s) (Connor et al., 2004). 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.  This biotope has a higher percentage of fine sediments than other Ascophyllum nodosum biotopes found on bedrock.  This increases the amount of sediment that could be entrained with higher water flow rates, which could increase the pressure from scour on the biotope at the sediment rock interface.  However, this is probably not relevant at the pressure benchmark.  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).

Sensitivity assessment.  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. However, an increase in the emergence time for a year is likely to change in the height of the biotope on the shore, with the top of the biotope being most sensitive to change as it is already at the upper tolerance limits. Conversely, a decrease in emergence may allow the biotope to increase its extent up the shore.  Overall, an increase in emergence is likely to see all of the biotopes on the shore shifting downwards. Ascophyllum nodosum can take as many as twelve years to recover, with the return of ecosystem function taking considerably longer. Therefore, the resistance of this biotope has been assessed as ‘Low’ and resilience as ‘Low’ so that sensitivity is assessed as ‘High’ to changes in emergence regime at the pressure benchmark.

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 20cm 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 a very to extremely sheltered biotope a further decrease in wave exposure is unlikely, and not significant given the very strong to strong tidal flow in which the biotope occurs. 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. For example, if the increase in wave exposure mobilized the substratum, the rolling of cobbles and pebbles or even boulders could remove the Ascophyllum and other fucoid cover.  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.  This biotope is found in a mid-eulittoral position and consequently, a proportion of time will be spent in the air where oxygen is not limited.  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 characterizing species Ascophyllum nodosum would 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.  Those immobile species such as the barnacle Semibalanus balanoides may experience some mortality.  However, barnacles can completely recolonize within three years (Bennell, 1981). The sheltered to extremely sheltered conditions that are characteristic of this biotope mean that water mixing is not very strong.  Therefore, water movement within this area will not reverse any oxygen depletion quickly, possibly exacerbating any negative effects. However, the biotope occurs in the mid-littoral so that emergence will mitigate the effects of hypoxic surface waters. Therefore, 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

The 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 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 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.

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. 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.

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 hard rock substrata (boulders, cobbles and pebbles) lying on sediment. The hard rock component is required for the settlement and growth of the dominant fucoid (Ascophyllum) canopy. Loss of the hard rock component 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

This biotope occurs on hard rock substrata (boulders, cobbles and pebbles) lying on sediment. The hard rock component is required for the settlement and growth of the dominant fucoid (Ascophyllum) canopy. A soft sedimentary habitat or mobile coarse sediments such as gravel or shingle would be unsuitable for these species.  Increased sediment instability would also be likely to reduce habitat suitability for littorinids.  In sites with mobile cobbles and boulders increased scour results in lower densities of Littorina spp. compared with other, local sites with stable substratum (Carlson et al., 2006).  A change to a sedimentary biotope without suitable attachment surfaces would lead to the development of a biological assemblage more typical of soft sediment habitats and the biotope would be lost. 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.  

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 biological community within this biotope occurs either on or within mixed sediment.  If the top 30 cm of this biotope were to be removed the dominant fucoid canopy and its associated community would be removed.  Therefore, resistance and resilience are assessed as  ‘Low’ and sensitivity to this pressure assessed as ‘High’.

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

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). 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.

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 mixed substrata found within this biotope could be subject to disturbance by penetrative gear but the impact of this pressure depends on the depth and footprint of the disturbance. Passing bottom gears have the potential to move or turn boulders, cobbles and pebbles (Picton & Goodwin, 2007). There is a lack of evidence to assess the pressure. However, sub-surface disturbance would also disrupt the surface layer of this biotope where the dominant fucoid (Ascophyllum) canopy is found. Therefore, the sensitivity assessment of this 'penetration' pressure is probably the same assessment for the 'abrasion/disturbance' pressure 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.

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 on the mid intertidal shore and consequently is subject to long periods of emersion during which time macroalgae can continue to photosynthesize as long as plants have sufficient water content. Therefore, photosynthesis and consequently growth will not be greatly affected. The level of water movement through wave exposure and tidal streams is unlikely to be high enough to cause any significant damage through scour.  Hence, resistance and resilience have been assessed as ‘High’. The sensitivity of this biotope to this pressure at the benchmark is assessed as ‘Not Sensitive’.

Smothering and siltation rate changes (light)

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

Evidence

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).

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 not excessive 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

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

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

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

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

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

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

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

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

In contrast to Didemnum vexillum’s preference for sheltered conditions, established colonies observed in Georges Bank and Long Island Sound were exposed to moderately strong tidal currents (1 to 2 knots; ca 0.5 to 1 m/s recorded at both sites) that may mobilise sediment (Valentine et al., 2007b; Mercer et al., 2009; Tillin et al., 2020). However, Valentine et al. (2007b) describe the substratum as immobile, presumably consolidated gravel, cobbles and pebbles.

Kleeman (2009), stated that the presence of a consistent mild wave action or ‘swash zone’ appears to favour Didemnum sp. establishment in the intertidal. Although some evidence suggests that waves and currents can facilitate the fragmentation and spread of Didemnum vexillum (Mckenzie et al., 2017), the tidal current velocities at some sites where Didemnum vexillum has been reported (for example, New England, where current velocities reach up to around 3 m/s) is lower than the current velocity required for the dislodgement of Didemnum vexillum fragments (around 7.6 m/s) (Reinhardt et al., 2012). This suggests that not all tidal currents are likely to dislodge Didemnum vexillum fragments. When on boat hulls the species can experience higher current velocities which is enough to cause dislodgement (Reinhardt et al., 2012).  

The Sandwich tide pools (USA) were subject to air exposure at low tide, and daily changes in water depth and temperatures (Valentine et al., 2007a). Didemnum vexillum colonies survived exposure to air at low tides for a short time (not exceeding two hours) during rapid colony growth in the summer months of July to September (Valentine et al., 2007a). However, parts of the large established colonies, which were artificially exposed to air for two to three hours in October, were observed desiccated or predated on by grazing periwinkles 30 days later, in the winter month of November (Valentine et al., 2007a). They suggested that the invasive tunicates’ ability to tolerate exposure to air varies with the seasonal growth cycle. Didemnum vexillum also tolerated emersion in Kent, as colonies on the mid-shore at Reculver flourish and survive in air exposure for up to three hours per cycle during springs (Hitchin, 2012). Hitchin (2012) suggested the porous nature of the sandstone boulders the species colonized retained water. The Kent shore was sheltered but held water due to its shallow slope and flats, which may allow Didemnum sp. to survive in the low to mid-shore. There is evidence that Didemnum vexillum died when exposed to air for more than six hours (Laing et al., 2010).

Undaria pinnatifida and Sargassum muticum  Thompson & Schiel (2012) found that native fucoids show high resistance to invasions by 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 has 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 in 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.

Didemnum vexillum requires hard substrata for successful colonization, therefore, it could colonize the rock and mixed sediment substratum typical of this biotope. Also, Didemnum vexillum has a preference for wave sheltered conditions. Didemnum vexillum has been recorded in the lower intertidal but in the mid-shore examples of the biotope, the abundance and extent of colonies may be limited due to emersion. Didemnum vexillum colonies can survive exposure to air at low tides for a short time (not exceeding two hours) (Valentine et al., 2007a). However, wave splash may mitigate Didemnum vexillum decline in the mid-shore by providing moisture. Didemnum vexillum can overgrow and displace sessile organisms, including brown algae species Ascophyllum nodosum, Sargassum, Fucus evanescens and Fucus serratus. There is no evidence on how Didemnum vexillum affects fucoids or if it causes fucoid mortality. However, Didemnum vexillum competing for light and space with fucoids and epifauna could lead to mortality and a reduction in biodiversity and may interfere with the recruitment of characteristic species. Therefore, a resistance of 'Medium' (some mortality, <25%) is suggested as a precaution. Resilience is likely to be 'Very low' as Didemnum vexillum would need to be physically removed to allow recovery. Hence, sensitivity to invasion by Didemnum is assessed as 'Medium'. 

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’.