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Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
Photographer: Christine Howson Copyright: Joint Nature Conservation Committee (JNCC)
Photographer: Sue Scott Copyright: Joint Nature Conservation Committee (JNCC)
Photographer: Anon. Copyright: Joint Nature Conservation Committee (JNCC)
Biotope distribution data provided by the JNCC (mhc.jncc.gov.uk)
Researched by
Thomas Stamp & Emma Williams
Refereed by
This information is not refereed
Thomas Stamp & Emma Williams Refereed by
UK and Ireland classification
EUNIS 2008
A3.2121 Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
JNCC 2022
IR.MIR.KR.LhypT.Ft Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
1997 Biotope
IR.MIR.KR.Lhyp.TFt Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
A3.2121 - Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
IR.MIR.KR.LhypT.Ft - Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
IR.MIR.KR.Lhyp.TFt - Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock
Moderately exposed, tide-swept, rock with dense Laminaria hyperborea forest is characterized by a rich under-storey and stipe flora of foliose seaweeds such as Plocamium cartilagineum, Metacallophyllis laciniata, Heterosiphonia plumosa, Cryptopleura ramosa and Delesseria sanguinea and crustose algae. Although these species are also found in most kelp forests, in this biotope they are particularly dense. On the rock surface, a rich fauna comprising sponges, anemones (such as Alcyonium digitatum, Cylista elegans and Urticina felina), hydroids, colonial ascidians and bryozoans. At some sites, instead of being covered by red algae, the kelp stipes may be heavily epiphytised by the sponge Halichondria panicea or the bryozoan Alcyonidium diaphanum. From some areas (such as Orkney), particularly good examples of this biotope may contain maerl and / or rhodoliths, with associated fauna between boulders. Both the flora and fauna of this biotope can be similar to the wave-exposed kelp forest (EIR.LhypFa) and although MIR.Lhyp.TFt is likely to contain species that are unable to tolerate strong wave action, further data analysis is required to clarify the differences. An example of this biotope was found in Lashy Sound in Orkney where Laminaria digitata dominated the tide-swept rock due to the very strong tides and/or non-laminar flow of water (Information from Connor et al., 2004; JNCC, 2015).
Depth range
0-5 m, 5-10 m, 10-20 m
UKBAP
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Sensitivity reviewHow is sensitivity assessed?
Sensitivity characteristics of the habitat and relevant characteristic species
At high densities, Laminaria hyperborea forms a canopy over infralittoral rock and mixed substrata. Beneath the canopy an understorey community grows, defined by a mixed red seaweed and faunal (filter feeding) turf. The abundance of Laminaria hyperborea is determined by light availability, which decreases with an increase in water depth. Therefore, depth and water clarity determine the density of Laminaria and hence the distribution of kelp forest (high density kelp) and park (low density kelp) variants. What distinguishes IR.MIR.KR.LhypT & IR.MIR.KR.LhypTX biotopes from other Laminaria hyperborea biotopes (e.g. IR.HIR.KFaR.LhypR) is exposure to strong (1.5-3 m/s)-moderately strong (0.5-1.5 m/s) tidal streams, which encourages the abundant growth of filter feeding fauna within the kelp understorey. Laminaria hyperborea stipes can be dominated by dense Botryllus schlosseri rather than the red seaweed communities of IR.HIR.KFaR.LhypR, and species such as Balanus crenatus and can be more abundant. Red seaweeds are also an important component of the understorey and stipe communities, however, in IR.MIR.KR.LhypT & IR.MIR.KR.LhypTX biotopes fauna are more predominant within the understorey than in other Laminaria hyperborea biotopes (except in IR.HIR.KFaR.LhypFa).
Kelp biotopes are a major source of primary productivity, and support magnified secondary productivity within North Atlantic coastal waters (Smale et al., 2013; Brodie et al., 2014). In Scotland alone, kelp biotopes are estimated to cover 8000km2 (Walker, 1953), and account for ca 45% of primary production in UK coastal waters (Smale et al., 2013). Therefore kelp biotopes, of which Laminaria hyperborea is dominant within UK sub-tidal rocky reefs (Birkett et al., 1998b), make a substantial contribution to coastal primary production in the UK (Smale et al., 2013). Laminaria hyperborea is grazed directly by species such as Patella pellucid, however approximately 80% of primary production is consumed as detritus or dissolved organic material (Krumhansl, 2012) which is both retained within and transported out of the parent kelp forest, providing valuable nutrition to potentially low productivity habitats such as sandy beaches (Smale et al., 2013).
Laminaria hyperborea acts as an ecosystem engineer (Jones et al., 1996; Smale et al., 2013) by altering; light levels (Sjøtun et al., 2006), physical disturbance (Connell, 2003), sedimentation rates (Eckman et al., 1989) and water flow (Smale et al., 2013), profoundly altering the physical environment for fauna and flora in close proximity. Laminaria hyperborea biotopes increase the three dimensional complexity of unvegetated rock (Norderhaug, 2004; Norderhaug et al., 2007; Norderhaug & Christie, 2011; Gorman et al., 2012; Smale et al., 2013), and support high local diversity, abundance, and biomass of epi/benthic species (Smale et al., 2013), and serve as a nursery ground for a number of commercially important species, e.g. Atlantic cod and pollack (Rinde et al., 1992).
In undertaking this assessment of sensitivity, an account is taken of knowledge of the biology of all characterizing species/taxa in the biotope. However, 'indicative species' are particularly important in undertaking the assessment because they have been subject to detailed research. For this sensitivity assessment, Laminaria hyperborea is the primary focus of research. However, it is recognized that the understorey community also defines the biotope. Examples of important species groups are mentioned where appropriate.
Resilience and recovery rates of habitat
A number of review and experimental publications have assessed the recovery of Laminaria hyperborea kelp beds and the associated community. If environmental conditions are favourable Laminaria hyperborea can recover following disturbance events reaching comparable plant densities and size to pristine Laminaria hyperborea beds within 2-6 years (Kain, 1979; Birkett et al., 1998b; Christie et al., 1998). Holdfast communities may recover in six years (Birkett et al., 1998b). Full epiphytic community and stipe habitat complexity regeneration required over 6 years (possibly 10 years). These recovery rates were based on discrete kelp harvesting events. Recurrent disturbance occurring frequently within 2-6 years of the initial disturbance is likely to lengthen recovery time (Birkett et al., 1998b; Burrows et al., 2014). Kain (1975) cleared sublittoral blocks of Laminaria hyperborea at different times of the year for several years. The first colonizers and succession community differed between blocks and at what time of year the blocks were cleared, however within 2 years of clearance the blocks were dominated by Laminaria hyperborea.
In south Norway, Laminaria hyperborea forests are harvested, which results in large scale removal of the canopy-forming kelps. Cristie et al. (1998) found that in south Norwegian Laminaria hyperborea beds a pool of small (<25 cm) understorey, Laminaria hyperborea plants persist beneath the kelp canopy for several years. The understorey Laminaria hyperborea sporophytes had fully re-established the canopy at a height of 1 m within 2-6 years after kelp harvesting. Within 1 year following harvesting, and each successive year thereafter, a pool of Laminaria hyperborea recruits had re-established within the understorey beneath the kelp canopy. Cristie et al. (1998) suggested that Laminaria hyperborea bed re-establishment from understorey recruits (see above) inhibits the colonization of other kelps species and furthers the dominance of Laminaria hyperborea within suitable habitats, stating that Laminaria hyperborea habitats are relatively resilient to disturbance events.
Laminaria hyperborea has a heteromorphic life strategy. A vast number of zoospores (mobile asexual spores) are released into the water column between October-April (Kain & Jones, 1964). Zoospores settle onto rock substrata and develop into dioecious gametophytes (Kain, 1979) which, following fertilization, develop into sporophytes and mature within 1-6 years (Kain, 1979; Fredriksen et al., 1995; Christie et al., 1998). Laminaria hyperborea zoospores have a recorded dispersal range of ca 200 m (Fredriksen et al., 1995). However zoospore dispersal is greatly influenced by water movements, and zoospore density and the rate of successful fertilization decreases exponentially with distance from the parental source (Fredriksen et al., 1995). Hence, recruitment following disturbance can be influenced by the proximity of mature kelp beds producing viable zoospores to the disturbed area. (Kain, 1979, Fredriksen et al., 1995).
Laminaria hyperborea biotopes are partially reliant on low (or no) populations of sea urchins, primarily the species; Echinus esculentus, Paracentrotus lividus and Strongylocentrotus droebachiensis, which graze directly on macroalgae, epiphytes, and the understorey community. Multiple authors (Steneck et al., 2002; Steneck et al., 2004; Rinde & Sjøtun, 2005; Norderhaug & Christie, 2009; Smale et al., 2013) have reported dense aggregations of sea urchins to be a principal threat to Laminaria hyperborea biotopes of the North Atlantic. Intense urchin grazing creates expansive areas known as 'urchin barrens', in which a shift can occur from Laminaria hyperborea dominated biotopes to those characterized by coralline encrusting algae, with a resultant reduction in biodiversity (Leinaas & Christie, 1996; Steneck et al., 2002, Norderhaug & Christie, 2009). Continued intensive urchin grazing pressure on Laminaria hyperborea biotopes can inhibit the Laminaria hyperborea recruitment (Sjøtun et al., 2006) and cause urchin barrens to persist for decades (Cristie et al., 1998; Stenneck et al., 2004; Rinde & Sjøtun, 2005). The mechanisms that control sea urchin aggregations are poorly understood but have been attributed to anthropogenic pressure on urchin predators (e.g. cod or lobsters). While these theories are largely unproven a number of studies have shown that removal of urchins from grazed areas coincides with kelp re-colonization (Leinaas & Christie, 1996; Norderhaug & Christie, 2009). Leinaas & Christie, (1996) removed Strongylocentrotus droebachiensis from 'urchin barrens' and observed a succession effect, in which the substratum was initially colonized by filamentous macroalgae and Saccharina latissima. However, after 2-4 years, Laminaria hyperborea dominated the community.
Reports of large scale urchin barrens within the North East Atlantic are generally limited to regions of the North Norwegian and Russian Coast (Rinde & Sjøtun, 2005, Norderhaug & Christie, 2009). Within the UK, urchin grazed biotopes (IR.MIR.KR.Lhyp.GzFt/Pk, IR.HIR.KFaR.LhypPar, IR.LIR.K.LhypSlat.Gz & IR.LIR.K.Slat.Gz) are generally localised to a few regions in North Scotland and Ireland (Smale et al., 2013; Stenneck et al., 2002; Norderhaug & Christie 2009; Connor et al., 2004). IR.MIR.KR.Lhyp.GzFt/Pk, IR.HIR.KFaR.LhypPar, IR.LIR.K.LhypSlat.Gz & IR.LIR.K.Slat.Gz are characterized by canopy-forming kelp. However, urchin grazing decreases the abundance and diversity of understorey species. In the Isle of Man. Jones & Kain (1967) observed that low Echinus esculentus grazing pressure could control the lower limit of Laminaria hyperborea and remove Laminaria hyperborea sporelings and juveniles. Urchin abundances in 'urchin barrens' have been reported as high as 100 individuals/m2 (Lang & Mann, 1976). Kain (1967) reported urchin abundances of 1-4/m2 within experimental plots of the Isle of Man. Therefore, while 'urchin barrens' are not presently an issue within the UK, relatively low urchin grazing has been found to control the depth distribution of Laminaria hyperborea, negatively impacting Laminaria hyperborea recruitment and reducing the understorey community abundance and diversity.
Other factors that are likely to influence the recovery of Laminaria hyperborea biotopes is competitive interactions with Invasive Non Indigenous Species (INIS), e.g. Undaria pinnatifida (Smale et al., 2013; Brodie et al., 2014; Heiser et al., 2014), and/or the Lusitanian kelp Laminaria ochroleuca (Brodie et al., 2014; Smale et al., 2014). A predicted sea temperature rise in the North and Celtic seas of between 1.5-5°C over the next century (Philippart et al., 2011) is likely to create northward range shifts in many macroalgal species, including Laminaria hyperborea. Laminaria hyperborea is a northern (Boreal) kelp species, thus increases in seawater temperature is likely to affect the resilience and recoverability of Laminaria hyperborea biotopes with southerly distributions in the UK (Smale et al., 2013; Stenneck et al., 2002). Evidence suggests that the Lusitanian kelp Laminaria ochroleuca (Smale et al., 2014), and the INIS Undaria pinnatifida (Heiser et al., 2014) are competing with Laminaria hyperborea along the UK south coast and may displace Laminaria hyperborea from some sub-tidal rocky reef habitats. The wider ecological consequences of Laminaria hyperborea' competition with Laminaria ochroleuca and Undaria pinnatifida are however as of yet unknown.
Resilience assessment. The evidence suggests that beds of mature Laminaria hyperborea can regenerate from disturbance within a period of 1-6 years, and the associated community within 7-10 years. However, other factors such as competitive interactions with Laminaria ochroleuca and Undaria pinnatifida may limit recovery of Laminaria hyperborea biotopes following disturbance. Similarly, changes in kelp canopy structure may alter sub-canopy environment conditions and therefore the understory algal assemblages could be influenced (Smale et al., 2020). Also, urchin grazing pressure is shown to limit Laminaria hyperborea recruitment and reduce the diversity and abundance of the understorey community and may limit habitat recovery following disturbance. The recovery of Laminaria hyperborea biotopes to disturbance from commercial harvesting in south Norway suggests that Laminaria hyperborea beds and the associated community could recover from a significant loss of canopy cover within 10 years, resilience has therefore been assessed as Medium. An exception is made for permanent or ongoing (long-term) pressures where recovery is not possible as the pressure is irreversible in which case resilience is assessed as 'Very Low' by default.
Please note in northern Norway urchin grazing pressure could extend recovery/resilience of the Laminaria hyperborea biotopes >25 years. If intensive urchin grazing (as seen in northern Norway) occurs in the UK resilience would be re-assessed as Very Low. However, because of the limited/localised incidence of urchin grazing within the UK, urchin grazing on large scales (as in Northern Norway) has not been included in this general resilience assessment. Introduction of Invasive Non-Indigenous Species (INIS) will also inhibit the recovery of Laminaria hyperborea biotopes for an indeterminate amount of time, in these cases, resilience would need to be re-assessed as Very Low. Another factor that is beyond the scope of this sensitivity assessment is the presence of multiple concurrent synergistic or cumulative effects, which Smale et al. (2013) suggest could be more damaging than the individual pressures.
Climate Change Pressures
Use / to open/close text displayed Resistance Resilience Sensitivity
Global warming (extreme)
Low Very Low High
Q: High
A: High
C: High
The distribution of kelp is strongly influenced by climatic conditions. Therefore, kelp species are extremely sensitive to the ongoing ocean warming (Kain, 1979; Van Den Hoek, 1982; Breeman, 1990; Lüning, 1990; Assis et al., 2016; Smale, 2020). Northern distribution boundaries are set by winter temperatures that are lethal, or summer temperatures too low for growth and/or reproduction, whilst southern limits are set by high lethal summer temperatures or winter temperatures too high for induction of a crucial step in the life cycle (Breeman, 1990). Kelps have a high dependence on ocean temperatures, which make them highly vulnerable to ocean warming (Assis et al., 2014). As temperatures increase, populations found towards the upper limit of their temperature range may be adversely affected by warming as physiological thresholds are exceeded (Wiens, 2016). Thermal stress can lead to mortality and consequent population-level effects, such as decreased abundance, altered size structure, local extinction and range contractions (Smale, 2020).
Laminaria hyperborea is a cold- temperate kelp species, distributed from the Barents Sea down to the coast of Portugal (Schoschina, 1997). Laminaria hyperborea has an optimum temperature for growth of 15°C, and an upper temperature limit of 21°C (Bolton & Lüning, 1982). At 17°C gamete survival is reduced (Steinhoff et al., 2008) and gametogenesis is inhibited at 21°C (Dieck, 1992). Therefore, Laminaria hyperborea recruitment could be impaired at a sustained temperature increase above 17°C. However, sporophytes can tolerate slightly higher temperatures of 20°C. Temperature tolerances for Laminaria hyperborea are also seasonally variable and temperature changes are less tolerated in winter months than summer months (Birkett et al., 1998b). Since 1970 Laminaria hyperborea has undergone a range constriction of ~250 km at its southern edge, with current persisting populations having reduced longevity and less reproductive individuals than those populations from past studies (Fernandez, 2011; Assis et al., 2016).
There is evidence that climate change is already having an impact on Laminaria hyperborea populations in the English Channel. Poleward range expansion of the warm temperate Laminaria ochroleuca as a result of ocean warming has led to competition with Laminaria hyperborea in UK waters (Smale et al., 2015). Laminaria ochroleuca was not found in the UK last century. But Laminaria ochroleuca has now increased its range to include the southwest of England (Smale et al., 2015) and the west coast of Ireland (Schoenrock et al., 2019).
During the 2013-2014 Northeast Atlantic storm season, the UK was subjected to some of the most intense storms recorded within the past five years. A study by Smale & Vance. (2015) investigated the impacts of the storms on kelp canopies along the south coast of the UK, findings indicated monospecific canopies of Laminaria hyperborea were unaffected by the storms. However, the storms significantly altered a mixed canopy study site, composed of Laminaria ochroleuca, Saccharina latissima and Laminaria hyperborea. Therefore, if climate change continues to change species composition within kelp forests resistance to storm disturbance could be altered.
Smale et al. (2015) found that Laminaria hyperborea suffered from much higher epiphytic loadings and lower productivity than its competitor Laminaria ochroleuca during the summer months, which reduced its competitive ability. The decreased competitive ability because of ocean warming corresponds to findings by Pessarrodona et al. (2018), who found a decrease in the size of Laminaria hyperborea plants along a north-south gradient in Scotland, with average maximum stipe lengths of over 150 cm, whereas in southern England they were less than 100 cm. Similarly, Smale et al. (2020b) observed clear differences between net primary productivity (NNP) and carbon standing stock of Laminaria hyperborea between the colder northern and warmer southern test sites in the UK, with NNP and standing stock being 1.5 and 2.5 times greater in the northern sites. Identifying ocean temperatures as a lively driver of productivity, with reduced NNP and standing stock observed in warmer waters (Smale et al., 2020b).
The decrease in productivity in southern England suggests that Laminaria hyperborea is already growing at suboptimal temperatures. Assis et al. (2018) predicted that under the highest emission scenario (RCP 8.5) the biogeographic range of Laminaria hyperborea will move northwards, and this retreat would lead to the species being lost from approximately 30% of the coastline of the UK.
Subtidal red algae are less tolerant of temperature extremes than intertidal red algae, surviving between -2°C and 18-23°C (Lüning 1990; Kain & Norton, 1990). Temperature increase may affect growth, recruitment, or interfere with reproduction processes. For example, there is some evidence to suggest that blade growth in Delesseria sanguinea is delayed until ambient sea temperatures fall below 13°C. Blade growth is also likely to be intrinsically linked to gametangia development (Kain, 1987), and maintenance of sea temperatures above 13°C may affect recruitment success.
Sensitivity assessment. Laminaria hyperborea is already growing at suboptimal temperatures in southern UK, based on evidence of decreased productivity comparative to Scotland (Pessarrodona et al., 2018; Smale et al., 2020b), and predictions have estimated Laminaria hyperborean to be lost from the UK by 2100 as a result of warming (Brodie et al., 2014). Sea surface temperatures around the UK currently range from between 6-19°C (Huthnance, 2010).
Under the middle emission scenario, a rise of 3°C could lead to maximum summer high temperatures in the south of the UK of 22°C. This is above the upper thermal limit of 21°C for Laminaria hyperborea (Bolton & Lüning, 1982), and is likely to lead to loss of this species from the south of England. Furthermore, biomass and plant size are expected to decrease as waters warm, with Scottish Laminaria hyperborea stipe lengths decreasing to lengths observed in southern England, leading to a decline in carbon assimilation, productivity and habitat quality. Therefore, resistance is assessed as 'Medium', and resilience is assessed as 'Very Low', as a loss is likely to be a long-term decline, due to the long-term nature of ocean warming. Therefore, this biotope is assessed as 'Medium' sensitivity to ocean warming under this scenario.
For the high emission scenario and extreme scenario, this northward retreat of the distribution of Laminaria hyperborea is expected to increase. Under the high emission scenario it is expected to be lost from 30% of the coastline around the UK (Assis et al., 2018), and under the extreme scenario even more is projected to be lost. Populations of Laminaria hyperborea that remain around the UK are predicted to become less productive. Therefore, under these scenarios, resistance is assessed as 'Low', and resilience is assessed as 'Very Low'. Therefore, this biotope is assessed as 'High' sensitivity to ocean warming under this scenario.
Global warming (high)
Global warming (middle)
Medium Very Low Medium
Marine heatwaves (high)
Low Low High
Q: Medium
A: Medium
Marine heatwaves are extreme weather events defined as periods of extreme sea surface temperature that persists for days to months (Frölicher et al., 2018). Marine heatwaves are predicted to occur more frequently, last for longer and at increased intensity by the end of this century under both middle and high emission scenarios (Frölicher et al., 2018). Marine heatwaves are known to cause significant impacts to kelp forests, particularly if a population is found towards the edge of its southern limit (Smale et al., 2019).
In Baja California, Mexico, an extreme heat even between 2014– 2016, led to both a decrease in density of Macrocystis pyrifera and a decrease in the number of fronds per individual in Baja California, Mexico (Arafeh-Dalmau et al., 2019). Additionally, there was a significant change to the understory algal composition, and half of the fish and invertebrates associated with this habitat disappeared. The same heatwave, coupled with a loss of starfish through disease and an increase in urchin grazing, led to the loss of > 90% of Macrocystis pyrifera from 350 km of coastline in northern California (Rogers-Bennett & Catton, 2019).
Under experimental conditions, Nepper-Davidson et al. (2019) exposed a northern (Denmark) population of Saccharina lattisima to a simulated three week heatwaves of three different intensities; 18, 21 and 24°C. When exposed to heatwaves of 18 and 21°C there was a decrease in photosynthesis and growth. When a 24°C was simulated, 91% of sporophytes were dead within a week, and the fronds of the few survivors were disintegrating, so the experiment was terminated (Nepper-Davidsen et al., 2019).
Laminaria hyperborea is a cold-temperate species of kelp with an optimum temperature for growth of 15°C, and an upper temperature limit of 21°C (Bolton & Lüning, 1982). Germination success can decrease by almost two thirds at temperatures as low as 17°C. Therefore, it is expected that, similar to other kelp species, Laminaria hyperborea will be highly sensitive to marine heatwaves.
Sensitivity assessment. Under the middle emission scenario, if heatwaves occurred every three years, with a maximum intensity of 2°C for 80 days by the end of this century, this could lead to summer sea temperatures reaching up to 24°C in southern England. Laminaria hyperborea is unlikely to survive a heatwave of this magnitude and is, therefore, likely to suffer severe mortality in the south. In Scotland, where a significant portion of Laminaria hyperborea populations occur, temperatures are not predicted to rise above 20°C, and therefore, Laminaria hyperborea is likely to survive a heatwave of this magnitude. Therefore, resistance has been assessed as 'Medium'. As a further heatwave (as defined by the pressure benchmark) is likely to affect this habitat before full recovery, resilience has been assessed as 'Low.' Therefore, this biotope is assessed as having 'Medium' sensitivity to marine heatwaves under the middle emission scenario.
Under the high emission scenario, if heatwaves occur every two years by the end of this century, reaching a maximum intensity of 3.5°C for 120 days, this could lead to the heatwave lasting the entire summer with temperatures reaching up to 26.5°C in southern England. Laminaria hyperborea is unlikely to survive a heatwave of this magnitude, and as temperatures are likely to reach > 21°C in Scotland under this scenario, there is likely to be mortality throughout this species' UK biogeographic distribution. Therefore, resistance has been assessed as 'Low'. As a further heatwave is likely to affect this habitat before full recovery (under the pressure benchmark definition), resilience has been assessed as 'Low.' Therefore, this biotope is assessed as having 'High' sensitivity to marine heatwaves under the high emission scenario.
Marine heatwaves (middle)
Medium Low Medium
Ocean acidification (high)
High High Not sensitive
C: Medium
Increasing levels of CO2 in the atmosphere have led to the average pH of sea surface waters dropping from 8.25 in the 1700s to 8.14 in the 1990s (Jacobson, 2005), with it expected to drop up to a further 0.35 units by the end of this century, dependent on emission scenario. Marine autotrophs will generally benefit from ocean acidification, through an increase in the availability of aqueous CO2 for photosynthesis (Koch et al., 2013).
Most species of kelp, including Laminaria hyperborea, appear to be undersaturated in respect to carbon dioxide, although they can generally utilise HCO3 and have external carbonic anhydrase for extracellular dehydration of HCO3 to CO2(Koch et al., 2013). This was confirmed for Laminaria hyperborea by Olischläger et al. (2012) who found that ocean acidification at levels expected for the end of this century (700 µatm CO2; a value between the middle and high emission scenario) led to an increase in female gametogenesis and increasing net photosynthesis and growth of sporophytes.
Research on other kelp species has revealed a positive or neutral effect of ocean acidification (Roleda et al., 2012, Fernández et al., 2015, Nunes et al., 2015, Iñiguez et al., 2016b, a), except for one study, which found that ocean acidification negatively impacted photosynthesis and growth in the southern hemisphere species, Ecklonia radiata (Britton et al., 2016).
Sensitivity assessment. Kelp forests live in a naturally variable pH habitat, with diel fluctuations of 0.3 - 0.45 pH units (Krause-Jensen et al., 2015, Britton et al., 2016), and boundary layer pH fluctuation of up to 0.8 units (Krause-Jensen et al., 2015). Laminaria hyperborea is not expected to be impacted by ocean acidification at levels expected for the end of this century. Therefore, under both the middle and high emission scenario resistance is assessed as 'High', and resilience is assessed as 'High' leading to a score of 'Not sensitive'.
Ocean acidification (middle)
Sea level rise (extreme)
Q: Low
A: NR
C: NR
A: Low
C: Low
Sea-level rise is occurring through a combination of thermal expansion and ice melt. Sea levels have risen 1-3 mm/yr in the last century (Cazenave & Nerem, 2004, Church et al., 2004, Church & White, 2006). This biotope IR.MIR.KR.LhypT.Ft occurs on extremely exposed, very exposed, exposed and moderately exposed, but wave surged, upper infralittoral bedrock and boulders. Wave surge is reduced with increasing depth; therefore, sea-level rise may reduce the effect of wave surge in the deeper examples or deeper reaches of this biotope.
The distribution of Laminaria hyperborea is positivity related to wave exposure, and as wave exposure increases, so does biomass and abundance (Pedersen et al., 2012). Total plant biomass and production per unit area double along an exposure gradient, due to larger leaf lengths and increased density at high wave energy sites (Pedersen et al., 2012). Understanding how sea-level rise will affect exposure and tidal energy is fraught with uncertainty, although evidence appears to suggest that any alterations will be non-linear (Pickering et al., 2012, Li et al., 2016). Modelling potential outcomes of sea-level rise on the tidal and residual currents in the Bohai Sea, China showed effects were site dependent, with energy either increasing or decreasing (Li et al., 2016). Similarly, Pickering et al. (2012) found a similar pattern around the UK for tidal amplitude.
Light availability and water turbidity are principle factors in determining kelp depth range (Birkett et al., 1998b), with laminarians being reported to be able to withstand light levels of up to 1% surface irradiance. An increase in depth due to sea-level rise is likely to impact both Laminaria hyperborea and any understory algae, negatively impacting this biotope. The most important factors explaining the distribution of Laminaria hyperborea along the Norwegian coast were depth, wave exposure, light exposure and topography (Bekkby et al., 2009). This species requires rocky substrate for attachment.
Sensitivity assessment. This biotope is recorded from 0 –20 m in depth. As wave surge diminishes with increased depth, sea-level rise is likely to lead to the density of faunal turf reducing at the deeper reaches of this biotope and transitioning into a biotope characterised by kelp and dense red seaweeds. Tidal streams are also likely to be reduced so that LhypT and its sub-biotopes may transition into kelp biotopes (IR.HIR.KFaR.LhypR.Ft or IR.HIR.KFaR.LhypR.Pk) with a lower faunal diversity or dense foliose red algae (IR.HIR.KFaR.FoR.Dic).LhpT.Pk and/or LhypTX.Pk made be lost as they represent the depth limit of Laminaria hyperborea. This biotope LhypT.Ft may be able to expand its range and migrate landwards to compensate for sea-level rise, if not constrained by lack of suitable substratum or human modified shorelines.
There is likely to be considerable variation between sites, the relative contribution of wave surge and exposure to habitat suitability, and the depth range occupied by the biotope. Hence, it is difficult to assess the effect of the different sea-level rise scenarios. However, as the biotope can occur from 0-20 m in depth, it is assumed at a sea-level rise of 0.5m, or 0.7 m (middle to high emission scenarios) would have limited effect but that a 1.07 m rise (the extreme emission scenario) might result in loss of some of the deeper extent of the biotope in some sites. Therefore, resistance is assessed as 'High' under the middle and high emission scenarios so that resilience is 'High' and sensitivity assessed as 'Not sensitive'. But resistance is possibly 'Medium' under the extreme emission scenario so that resilience is 'Very low' and sensitivity assessed as 'Medium', albeit with 'Low' confidence.
Sea level rise (high)
Sea level rise (middle)
Hydrological Pressures
Temperature increase (local)
Medium Medium Medium
Kain (1964) stated that Laminaria hyperborea sporophyte growth and reproduction could occur within a temperature range of 0 - 20°C. Upper and lower lethal temperatures have been estimated at between 1-2 °C above or below the extremes of this range (Birkett et al., 1988). Above 17°C gamete survival is reduced (Kain, 1964 & 1971) and gametogenesis is inhibited at 21 °C (Dieck, 1992). It is therefore likely that Laminaria hyperborea recruitment will be impaired at a sustained temperature increase of above 17°C. Sporophytes however can tolerate slightly higher temperatures of 20°C. Temperature tolerances for Laminaria hyperborea are also seasonally variable and temperature changes are less tolerated in winter months than summer months (Birkett et al., 1998).
Subtidal red algae are less tolerant of temperature extremes than intertidal red algae, surviving between -2°C and 18-23 °C (Lüning 1990; Kain & Norton, 1990). Temperature increase may affect growth, recruitment or interfere with reproduction processes. For example, there is some evidence to suggest that blade growth in Delesseria sanguinea is delayed until ambient sea temperatures fall below 13 °C. Blade growth is also likely to be intrinsically linked to gametangia development (Kain, 1987), and maintenance of sea temperatures above 13 °C may affect recruitment success.
Laminaria hyperborea has a geographic range from mid Portugal to Northern Norway (Birket et al., 1998), and a mid range within southern Norway (60°-65° North)(Kain, 1971). The average seawater temperature for southern Norway in October is 12-13°C (Miller et al., 2009), and average annual sea temperature, from 1970-2014, is 8°C (Beszczynska-Möller & Dye, 2013). Against the pressure benchmark the available information suggests that Laminaria hyperborea recruitment processes may be affected and associated red algae communities may decline.
Sensitivity assessment. Overall, a chronic change (2°C for a year) outside normal range for a year may reduce recruitment and growth, resulting in a minor loss in the population of kelp, especially in winter months or in southern examples of the biotope. However, an acute change (5°C for a month; e.g. from thermal effluent) may result in loss of abundance of kelp or extent of the bed, especially in winter. Therefore, resistance to the pressure is considered 'Medium', and resilience 'Medium'. The sensitivity of this biotope to increases in temperature has been assessed as 'Medium'.
Temperature decrease (local)
Kain (1964) stated that Laminaria hyperborea sporophyte growth and reproduction could occur within a temperature range of 0 - 20°C. Upper and lower lethal temperatures have been estimated at between 1-2 °C above or below the extremes of this range (Birkett et al., 1988). Subtidal red algae can survive at temperatures between -2 °C and 18-23 °C (Lüning, 1990; Kain & Norton, 1990).
Laminaria hyperborea is a boreal northern species with a geographic range from mid Portugal to Northern Norway (Birket et al., 1998), and a mid range within southern Norway (60°-65° North)(Kain, 1971). The average seawater temperature for southern Norway in October is 12-13°C (Miller et al., 2009), and average annual sea temperature, from 1970-2014, is 8°C (Beszczynska-Möller & Dye, 2013). The available information suggests that Laminaria hyperborea and biotope structure would not be affected by a change in sea temperature at the benchmark level.
Sensitivity assessment. Resistance to the pressure is considered 'High', and resilience 'High'. The sensitivity of this biotope to decreases in temperature has been assessed as 'Not Sensitive'.
Salinity increase (local)
Low Medium Medium
Lüning (1990) suggest that "kelps" are stenohaline, their general tolerance to salinity as a phenotypic group covering 16 - 50 psu over a 24 hr period. Optimal growth probably occurs between 30-35 psu (MNCR category- 'Full' salinity) and growth rates are likely to be affected by periodic salinity stress. Birkett et al, (1998) suggested that long-term increases in salinity may affect Laminaria hyperborea growth and may result in loss of affected kelp, and therefore loss of the biotope.
Sensitivity assessment. Resistance to the pressure is considered 'Low', and resilience 'Medium'. The sensitivity of this biotope to an increase in salinity has been assessed as 'Medium'.
Salinity decrease (local)
Lüning (1990) suggest that 'kelps' are stenohaline, their general tolerance to salinity as a phenotypic group covering 16 - 50 psu over a 24 hr period. Optimal growth probably occurs between 30-35 psu (MNCR category-Full Salinity) and growth rates are likely to be affected by periodic salinity stress. Birkett et al,. (1998) suggest that long-term changes in salinity may result in loss of affected kelp and, therefore loss of this biotope.
Hopkin & Kain (1978) tested Laminaria hyperborea sporophyte growth at various low salinity treatments. The results showed that Laminaria hyperborea sporophytes could grow 'normally' at 19 psu, growth was reduced at 16 psu and did not grow at 7 psu. A decrease in one MNCR salinity scale from Full Salinity (30-40psu) to Reduced Salinity (18-30 psu) would result in a decrease of Laminaria hyperborea sporophyte growth. Laminaria hyperborea may also be out-competed by low salinity tolerant species e.g. Saccharina latissma (Karsten, 2007), or invasive kelp species, e.g. Undaria pinnatifida (Burrows et al., 2014).
If salinity was returned to Full Salinity (30-40 psu) Laminaria hyperborea could out-compete Saccharina latissma and re-establish community dominance in 2-4 years (Leinaas & Christie, 1996), however full habitat structure may take over 10 years to recover (Birkett et al., 1998; Cristie et al., 1998). The ability of Laminaria hyperborea to out-compete Undaria pinnatifida within the UK is however unknown (Heiser et al., 2014), and as such interspecific interaction between Laminaria hyperborea and Undaria pinnatifida is not included within this sensitivity assessment.
Sensitivity assessment. Resistance to the pressure is considered 'Low', and resilience 'Medium'. The sensitivity of this biotope to decreases in salinity has been assessed as 'Medium'.
Water flow (tidal current) changes (local)
Kregting et al. (2013) measured Laminaria hyperborea blade growth and stipe elongation from an exposed and a sheltered site in Strangford Lough, Ireland, from March 2009-April 2010. Maximal significant wave height (Hm0) was 3.67 & 2m at the exposed and sheltered sites, and maximal water velocity (Velrms) was 0.6 & 0.3m/s at the exposed and sheltered sites respectively. Despite the differences in wave exposure and water velocity there was no significant difference in Laminaria hyperborea growth between the exposed and sheltered sites. Therefore water flow was found to have no significant effect on Laminaria hyperborea growth at the observed range of water velocities.
Biotope structure is however different between wave exposed and sheltered sites. Pedersen et al. (2012) observed Laminaria hyperborea biomass, productivity and density increased with an increase in wave exposure. At low wave exposure Laminaria hyperborea canopy forming plants were smaller, had lower densities and had higher mortality rates than at exposed sites. At low wave exposure Pedersen et al. (2012) suggested that high epiphytic loading on Laminaria hyperborea impaired light conditions, nutrient uptake, and increased the drag on the host Laminaria hyperborea during extreme storm events.
The morphology of the stipe and blade of kelps vary with water flow. In wave exposed areas, for example, Laminaria hyperborea develops a long and flexible stipe and this is probably a functional adaptation to strong water movement (Sjøtun, 1998). In addition, the lamina becomes narrower and thinner in strong currents (Sjøtun & Fredriksen, 1995). However, the stipe of Laminaria hyperborea is relatively stiff and can snap in strong currents. Laminaria hyperborea is usually absent from areas of high wave action or strong currents, although it is found in the Menai Strait, Wales, where tidal velocities can exceed 4 m/s (NBN, 2015) and in tidal rapids in Norway (J. Jones, pers. comm.) Laminaria hyperborea growth can persist in very strong tidal streams (>3 m/s).
Increase water flow rate may also remove or inhibit grazers including Patella pellucida and Echinus esculentus, therefore reducing grazing in the understorey and on stipes. The associated algal flora and suspension feeding faunal populations change significantly with different water flow regimes. Decreased water flow rates may reduce the suspension feeding understorey epifauna, to be replaced by an epiflora dominated community as in the biotope IR.HIR.KFaR.LhypR. The composition of the holdfast fauna may also change, e.g. energetic or sheltered water movements favour different species of amphipods (Moore, 1985).
IR.MIR.KR.LhypT, IR.MIR.KR.LhypTX and their associated sub-biotopes are predominantly found within strong (1.5-3 m/s)-moderate (0.5-1.5 m/s) tidal streams. The prominent understorey filter feeding community within IR.MIR.KR.LhypT/TX is reliant on strong tidal flow. It is the abundance of filter feeding organisms that separates the tide swept Laminaria hyperborea biotopes from the not tideswept biotopes within the same wave exposure (e.g. IR.HIR.KFaR.LhypR). A change in peak mean spring bed flow velocity within the tidal streams 0.5-3 m/s is not likely to significantly affect the abundance of Laminaria hyperborea. A decrease in tidal streams may result in a decline of filter feeding fauna and an increase in red seaweeds within the understorey community. A decrease in tidal flow within this range may also decrease urchin dislodgment and increase urchin grazing. An increase in urchin grazing may cause a decline in the understorey community abundance and diversity (as in IR.MIR.KR.Lhyp.GzFt/Pk). Large increases in water movement (e.g. >3 m/s) may increase the dislodgement/loss of Laminaria hyperborea from the biotope (Birkett et al., 1988), and may cause an increase in the abundance of the ephemeral kelp: Saccharina latissima or Alaria esculenta, which are both fast growing species and tolerant of fast water movement (Birket et al., 1998).
Sensitivity assessment. Water movement is a key environmental characteristic of IR.MIR.KR.LhypT and IR.MIR.KR.LhypTX, however these biotopes are found within a broad range of tidal streams (0.5-3 m/s). A change in peak mean spring bed flow velocity of between 0.1m/s to 0.2m/s for more than 1 year is however observed as a small change in water movement, and is not likely to significantly affect the community Resistance to the pressure is therefore considered 'High', and resilience 'High'. The sensitivity of this biotope to changes in peak mean spring bed velocity has been assessed as 'Not Sensitive' at the bench mark level.
Large and dramatic changes in tidal streams (>3m/sec) may increase the abundance/dominance of ephemeral kelp species Alaria esculenta and Saccharina latissima, and may result in loss of the biotope. Changes of this dramatic nature are however outside of the scope of this habitat sensitivity assessment.
Emergence regime changes
The upper limit of the Laminaria hyperborea bed is determined by wave action and water flow, desiccation, and competition from the more emergence resistant Laminaria digitata. Laminaria hyperborea exposed at extreme low water are very intolerant of desiccation, the most noticeable effect being bleaching of the frond and subsequent death of the meristem and loss of the plant. An increase in wave exposure (See below- water flow), as a result of increased emergence, has been found to exclude Laminaria hyperborea from shallow waters due to dislodgement of the sporophyte or snapping of the stipe (Birket et al., 1998). Hence, an increase in emergence is likely to lead to mortality of exposed Laminaria hyperborea and the associated habitat.
An increase in water depth/decreased emergence (at the benchmark level) may increase the upper depth restriction of Laminaria hyperborea forest variants within this biotope group. However, limited light availability at depth will decrease the lower extent of Laminaria hyperborea, and may therefore result in a shift from forest to park biotope variants at depth. Further increases in depth will cause a community shift to that characterized by circalittoral faunal species, however this is beyond the scope of the benchmark.
Sensitivity assessment. Resistance to the pressure is considered 'Low', and resilience 'Medium'. The sensitivity of this biotope to changes in tidal emergence has been assessed as 'Medium'.
Wave exposure changes (local)
Kregting et al. (2013) measured Laminaria hyperborea blade growth and stipe elongation from an exposed and a sheltered site in Strangford Lough, Ireland from March 2009-April 2010. Wave exposure was found to be between 1.1. to 1.6 times greater between the exposed and sheltered sites. Maximal significant wave height (Hm0) was 3.67 & 2 m at the exposed and sheltered sites. Maximal water velocity (Velrms) was 0.6 & 0.3 m/s at the exposed and sheltered sites. Despite the differences in wave exposure and water velocity there was no significant difference in Laminaria hyperborea growth between the exposed and sheltered site.
Biotope structure is however different between wave exposed and sheltered sites. Pedersen et al. (2012) observed Laminaria hyperborea biomass, productivity and density increased with an increase in wave exposure. At low wave exposure Laminaria hyperborea canopy forming plants were smaller, had lower densities and had higher mortality rates than at exposed sites. At low wave exposure high epiphytic loading on Laminaria hyperborea was theorised to impair light conditions, nutrient uptake, and increase the drag of the host Laminaria hyperborea during extreme storm events.
The morphology of the stipe and blade of kelps vary with water flow. In wave exposed areas, for example, Laminaria hyperborea develops a long and flexible stipe and this is probably a functional adaptation to strong water movement (Sjøtun, 1998). In addition, the lamina becomes narrower and thinner in strong currents (Sjøtun & Fredriksen, 1995). However, the stipe of Laminaria hyperborea is relatively stiff and can snap in strong currents. Lamiaria hyperborea is usually absent from areas of extreme wave action and can be replaced by Alaria esculenta. In extreme wave exposures Alaria esculenta can dominate the shallow sub-littoral to a depth of 15m, where Laminaria hyperboea dominates the infralittoral (Birket et al., 1998).
Increase water flow may also remove or inhibit grazers including Patella pellucida and Echinus esculentus, therefore reducing grazing in the understorey and on stipes. The associated algal flora and suspension feeding faunal populations change significantly with different water flow regimes. Increased water flow rates may reduce the understorey epiflora, to be replaced by an epifauna dominated community (e.g. sponges, anemones and polyclinid ascidians) as in the biotope IR.HIR.KFaR.LhypFa. The composition of the holdfast fauna may also change, e.g. energetic or sheltered water movements favour different species of amphipods (Moore, 1985).
IR.MIR.KR.LhypT, IR.MIR.KR.LhypTX and their associated sub-biotopes are found between extremely exposed to sheltered wave exposure but experience elevated tidal streams. Changes in local wave height above or below that experienced in extremely exposed to sheltered exposed sites will affect the dominance of Laminaria hyperborea. Smaller changes in local wave height have the potential to cause changes to the understorey community. The prominent understorey filter feeding community within IR.MIR.KR.LhypT/TX is reliant on high water movement. A decrease in wave surge may result in a decline of filter feeding fauna and an increase in red seaweeds within the understorey community or vice versa. A decrease in local wave height may also decrease the chance of urchins being dislodged (removed) from biotopes found at sites with traditionally high wave exposure and may therefore increase urchin grazing. An increase in urchin grazing may cause a decline in the understorey community abundance and diversity (as in IR.MIR.KR.Lhyp.GzFt/Pk and IR.MIR.KR.LhypPar).
Sensitivity assessment. A change in nearshore significant wave height >3% but <5% is however unlikely to have a significant effect. Resistance to the pressure is considered 'High', and resilience 'High'. Hence, the sensitivity of this biotope to changes in local wave height has been assessed as 'Not Sensitive'.
Large and dramatic changes in near shore wave height may increase the abundance/dominance of the ephemeral kelp species Alaria esculenta, increase the dominance of IR.HIR.KFaR.Ala and may result in loss of the biotope. Changes of this dramatic nature are however outside of the scope of this habitat sensitivity assessment.
Chemical Pressures
Transition elements & organo-metal contamination
Not Assessed (NA) Not assessed (NA) Not assessed (NA)
Q: NR
Bryan (1984) suggested that the general order for heavy metal toxicity in seaweeds is: Organic Hg > inorganic Hg > Cu > Ag > Zn > Cd > Pb. Cole et a,. (1999) reported that Hg was very toxic to macrophytes. Similarly, Hopkin & Kain (1978) demonstrated sub-lethal effects of heavy metals on Laminaria hyperborea gametophytes and sporophytes, including reduced growth and respiration. Sheppard et al., (1980) noted that increasing levels of heavy metal contamination along the west coast of Britain reduced species number and richness in holdfast fauna, except for suspension feeders which became increasingly dominant. Gastropods may be relatively tolerant of heavy metal pollution (Bryan, 1984). Echinus esculentus recruitment is likely to be impaired by heavy metal contamination due to the intolerance of its larvae. Echinus esculentus are long-lived and poor recruitment may not reduce grazing pressure in the short-term. Although macroalgae species may not be killed, except by high levels of contamination, reduced growth rates may impair the ability of the biotope to recover from other environmental disturbances.
This pressure is Not assessed but evidence is presented where available
Hydrocarbon & PAH contamination
This pressure is Not assessed but evidence is presented where available.
Laminaria hyperborea fronds, being almost exclusively sub tidal, would not come into contact with freshly released oil, but only to sinking emulsified oil and oil adsorbed onto particles (Birket et al., 1998). The mucilaginous slime layer coating of laminarians may protect them from smothering by oil. Hydrocarbons in solution reduce photosynthesis and may be algicidal. However, Holt et al,. (1995) reported that oil spills in the USA and from the 'Torrey Canyon' had little effect on kelp forests. Similarly, surveys of subtidal communities at a number sites between 1-22.5m below chart datum, including Laminaria hyperbora communities, showed no noticeable impacts of the Sea Empress oil spill and clean up (Rostron & Bunker, 1997). An assessment of holdfast fauna in Laminaria showed that although species richness and diversity decreased with increasing proximity to the Sea Empress oil spill, overall the holdfasts contained a reasonably rich and diverse fauna, even though oil was present in most samples (Sommerfield & Warwick, 1999). Laboratory studies of the effects of oil and dispersants on several red algae species, including Delesseria sanguinea (Grandy 1984; cited in Holt et al., 1995) concluded that they were all sensitive to oil/ dispersant mixtures, with little differences between adults, sporelings, diploid or haploid life stages. Holt et al., (1995) concluded that Delesseria sanguinea is probably generally sensitive of chemical contamination. Overall the red algae are likely to be highly intolerant to hydrocarbon contamination. Loss of red algae is likely to reduce the species richness and diversity of the biotope and the understorey may become dominated by encrusting corallines; however, red algae are likely to recover relatively quickly.
Synthetic compound contamination
O'Brian & Dixon (1976) suggested that red algae were the most sensitive group of macrophytes to oil and dispersant contamination (see Smith, 1968). Although Laminaria hyperborea sporelings and gametophytes are intolerant of atrazine (and probably other herbicides) overall they may be relatively tolerant of synthetic chemicals (Holt et al., 1995). Laminaria hyperborea survived within >55m from the acidified halogenated effluent discharge polluting Amlwch Bay, Anglesey, albeit at low density. These specimens were greater than 5 years of age, suggesting that spores and/or early stages were more intolerant (Hoare & Hiscock, 1974). Patella pellucida was excluded from Amlwch Bay by the pollution and the species richness of the holdfast fauna decreased with proximity to the effluent discharge; amphipods were particularly intolerant although polychaetes were the least affected (Hoare & Hiscock, 1974). The richness of epifauna/flora decreased near the source of the effluent and epiphytes were absent from Laminaria hyperborea stipes within Amlwch Bay. The red alga Phyllophora membranifolia was also tolerant of the effluent in Amlwch Bay. Smith (1968) also noted that epiphytic and benthic red algae were intolerant of dispersant or oil contamination due to the Torrey Canyon oil spill; only the epiphytes Crytopleura ramosa and Spermothamnion repens and some tufts of Jania rubens survived together with Osmundea pinnatifida, Gigartina pistillata and Phyllophora crispa from the sublittoral fringe. Delesseria sanguinea was probably to most intolerant since it was damaged at depths of 6m (Smith, 1968). Holt et al., (1995) suggested that Delesseria sanguinea is probably generally sensitive of chemical contamination. Although Laminaria hyperborea may be relatively insensitive to synthetic chemical pollution, evidence suggests that grazing gastropods, amphipods and red algae are sensitive. Loss of red algae is likely to reduce the species richness and diversity of the biotope and the understorey may become dominated by encrusting corallines; however, red algae are likely to recover relatively quickly.
Radionuclide contamination
Not relevant (NR) Not relevant (NR) No evidence (NEv)
No evidence
Introduction of other substances
This pressure is Not assessed.
De-oxygenation
Medium High Low
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). In addition, the biotope occurs in areas of moderate to extreme wave action, so is likely to be continuously aerated. A rapid recovery from a state of low oxygen is expected if the environmental conditions are transient. If levels do drop below 4 mg/l negative effects on these organisms can be expected with adverse effects occurring below 2mg/l (Cole et al., 1999).
Sensitivity Assessment. Reduced oxygen levels are likely to inhibit photosynthesis and respiration but not cause a loss of the macroalgae population directly. Furthermore wave exposure is likely to constantly aerate the affected area. While de-oxygenation may not directly affect Laminaria hyperborea, small invertebrate epifauna may be lost, causing a reduction in species richness. Therefore resistance has been assessed as 'Medium' is recorded. Resilience is likely to be 'High', and the biotopes is probably 'Low' at the benchmark level.
Nutrient enrichment
Not relevant (NR) Not relevant (NR) Not sensitive
Holt et al,. (1995) suggest that Laminaria hyperborea may be tolerant of nutrient enrichment since healthy populations are found at ends of sublittoral untreated sewage outfalls in the Isle of Man. Increased nutrient levels e.g. from sewage outfalls, has been associated with increases in abundance, primary biomass and Laminaria hyperborea stipe production but with concomitant decreases in species numbers and diversity (Fletcher, 1996).
Increased nutrients may result in phytoplankton blooms that increase turbidity (see above). Increased nutrients may favour sea urchins, e.g. Echinus esculentus, due their ability to absorb dissolved organics, and result in increased grazing pressure leading to loss of understorey epiflora/fauna, decreased kelp recruitment and possibly 'urchin barrens'. Therefore, although nutrients may not affect kelps directly, indirect effects such as turbidity, siltation and competition may significantly affect the structure of the biotope.
However this biotope is considered to be 'Not sensitive' at the pressure benchmark, that assumes compliance with good status as defined by the WFD.
Organic enrichment
Holt et al., (1995) suggest that Laminaria hyperborea may be tolerant of organic enrichment since healthy populations are found at ends of sublittoral untreated sewage outfalls in the Isle of Man. Increased nutrient levels e.g. from sewage outfalls, has been associated with increases in abundance, primary biomass and Laminaria hyperborea stipe production, but with concomitant decreases in species numbers and diversity (Fletcher, 1996). Increase organic enrichment has also been found to increase the abundance and dominance of suspension feeding fauna within Laminaria hyperborea holdfasts (Sheppard et al., 1980). Increase in ephemeral and opportunistic algae are associated with reduced numbers of perennial macrophytes (Fletcher, 1996). Increased nutrients may also result in phytoplankton blooms that increase turbidity. Therefore, although nutrients may not affect kelps directly, indirect effects such as turbidity and the increased abundance of suspension feeding fauna may affect the structure of Laminaria hyperborea biotopes (see water clarity above).
Sensitivity assessment. While organic enrichment may not have any direct effects on Laminaria hyperborea, increased turbidity and abundance of suspension feeding fauna may have significant effects on the biotope structure. Resistance to the pressure has therefore been considered 'Medium', and resilience 'High'. The sensitivity of this biotope to organic enrichment is assessed as 'Low'.
Physical Pressures
Physical loss (to land or freshwater habitat)
None Very Low High
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)
If rock substrata were replaced with sedimentary substrata this would represent a fundamental change in habitat type, which Laminaria hyperborea would not be able to tolerate (Birket et al., 1998). The biotope would be lost.
Sensitivity assessment. Resistance to the pressure is considered 'None', and resilience 'Very Low' or 'None'. The sensitivity of this biotope to change from sedimentary or soft rock substrata to hard rock or artificial substrata or vice-versa is assessed as 'High'.
Physical change (to another sediment type)
Not relevant (NR) Not relevant (NR) Not relevant (NR)
Habitat structure changes - removal of substratum (extraction)
None Medium Medium
IR.MIR.KR.LhypTX plus associated sub biotopes, are found on tide swept boulders, cobbles, pebbles and gravel. Extraction of the substratum would likely result in high mortality of Laminaria hyperborea plus the associated community.
Sensitivity assessment. Resistance has been assessed as 'None', resilience as 'Medium'. Sensitivity has been assessed as 'Medium'
Abrasion/disturbance of the surface of the substratum or seabed
Christie et al. (1998) observed Laminaria hyperborea habitat regeneration following commercial Laminaria hyperborea trawling in south Norway. Within the study area, trawling removed all large canopy-forming adult Laminaria hyperborea, however sub-canopy recruits were largely unaffected. In 2-6 years of harvesting a new canopy had formed 1 m off the seabed. The associated holdfast communities recovered in 6 years, however the epiphytic stipe community did not fully recover within the same time period. Christie et al. (1998) suggested that kelp habitats were relatively resistant to direct disturbance/removal of Laminaria hyperborea canopy.
Recurrent disturbance occurring at a smaller time scale than the recovery period of 2-6 years (stated above) could extend recovery time. Kain (1975) cleared sublittoral blocks of Laminaria hyperborea at different times of the year for several years. The first colonizers and succession community differed between blocks and at what time of year the blocks were cleared however within 2 years of clearance the blocks were dominated by Laminaria hyperborea. Leinaas & Christie (1996) also observed Laminaria hyperborea re-colonization of "urchin barrens", following removal of urchins. The substratum was initially colonized by filamentous macroalgae and Saccharina latissima however after 2-4 years Laminaria hyperborea dominated the community.
Sensitivity assessment. Resistance to the pressure is considered 'Low', and resilience 'Medium'. The sensitivity of this biotope to damage to seabed surface features is assessed as 'Medium'.
Penetration or disturbance of the substratum subsurface
Not relevant, please refer to pressure 'Abrasion/ disturbance of the substratum on the surface of the seabed'.
Changes in suspended solids (water clarity)
Suspended Particle Matter (SPM) concentration has a linear relationship with sub-surface light attenuation (Kd) (Devlin et al., 2008). An increase in SPM results in a decrease in sub-surface light attenuation. Light availability and water turbidity are principal factors in determining kelp depth range (Birkett et al., 1998). Light penetration influences the maximum depth at which kelp species can grow and it has been reported that laminarians grow down to depths at which the light levels are reduced to 1 percent of incident light at the surface. Maximal depth distribution of laminarians, therefore, varies from 100 m in the Mediterranean to only 6-7 m in the silt-laden German Bight. In Atlantic European waters, the depth limit is typically 35 m. In very turbid waters the depth at which Laminaria hyperborea is found may be reduced, or in some cases excluded completely (e.g. Severn Estuary), because of the alteration in light attenuation by suspended sediment (Birkett et al. 1998b; Lüning, 1990).
Laminaria spp. show a decrease of 50% photosynthetic activity when turbidity increases by 0.1/m (light attenuation coefficient =0.1-0.2/m; Staehr & Wernberg, 2009). An increase in water turbidity will likely affect the photosynthetic ability of Laminaria hyperborea and Laminaria ochroleuca and decrease Laminaria hyperborea abundance and density (see sub-biotope IR.MIR.KR.LhypT.Pk). Kain (1964) suggested that early Laminaria hyperborea gametophyte development could occur in the absence of light. Furthermore, observations from south Norway found that a pool of Laminaria hyperborea recruits could persist growing beneath Laminaria hyperborea canopies for several years, indicating that sporophyte growth can occur in light-limited environments (Christe et al., 1998). However in habitats exposed to high levels of suspended silts Laminaria hyperborea is out-competed by Saccharina latissima, a silt tolerant species, and thus, a decrease in water clarity is likely to decrease the abundance of Laminaria hyperborea in the affected area (Norton, 1978).
Sensitivity Assessment. Changes in water clarity are likely to affect photosynthetic rates and enable Saccharina latissima to compete more successfully with Laminaria hyperborea. A decrease in turbidity is likely to support enhanced growth (and possible habitat expansion) and is therefore not considered in this assessment. An increase in water clarity from clear to intermediate (10-100 mg/l) represents a change in light attenuation of ca 0.67-6.7 Kd/m, and is likely to result in a greater than 50% reduction in photosynthesis of Laminaria spp. Therefore, the dominant kelp species will probably suffer a significant decline and resistance to this pressure is assessed as 'Low'. Resilience to this pressure is probably 'Medium' at the benchmark. Hence, this biotope is assessed as having a sensitivity of 'Medium 'to this pressure.
Smothering and siltation rate changes (light)
Smothering by sediment e.g. 5 cm material during a discrete event, is unlikely to damage Laminaria hyperborea plants but is likely to affect gametophyte survival as well as holdfast fauna, and interfere with zoospore settlement. Given the microscopic size of the gametophyte, 5 cm of sediment could be expected to significantly inhibit growth. However, laboratory studies showed that gametophytes can survive in darkness for between 6 - 16 months at 8 °C and would probably survive smothering by a discrete event. Once returned to normal conditions the gametophytes resumed growth or maturation within 1 month (Dieck, 1993). Intolerance to this factor is likely to be higher during the peak periods of sporulation and/or spore settlement.
If inundation is long lasting then the understorey epifauna/flora may be adversely affected, e.g. suspension or filter feeding fauna and/or algal species. If clearance of deposited sediment occurs rapidly then understorey communities are expected to recover quickly. IR.MIR.KR.LhypT/TX (and their associated sub-biotopes) occur in high to moderate energy habitats (due to water flow or wave action) so deposited sediment is unlikely to remain for more than a few tidal cycles, except in the deepest of rock-pools.
Sensitivity assessment. Due to the strong tidal flows that characterize IR.MIR.KR.LhypT, IR.MIR.KR.LhypTX, deposited sediments are likely to be rapidly dispersed from the affected site. Resistance to the pressure is therefore considered 'High', and resilience 'High'. The sensitivity of this biotope to light deposition of up to 5cm of fine material added to the seabed in a single discreet event is assessed as 'Not Sensitive'.
Smothering and siltation rate changes (heavy)
Smothering by sediment e.g. 30 cm material during a discrete event, is unlikely to damage Laminaria hyperborea plants but is likely to affect gametophyte survival, holdfast communities, epiphytic community at the base of the stype, and interfere with zoospore settlement. Given the microscopic size of the gametophyte, 30 cm of sediment could be expected to significantly inhibit growth. However, laboratory studies showed that gametophytes can survive in darkness for between 6 - 16 months at 8 °C and would probably survive smothering within a discrete event. Once returned to normal conditions the gametophytes resumed growth or maturation within 1 month (Dieck, 1993). Intolerance to this factor is likely to be higher during the peak periods of sporulation and/or spore settlement.
Sensitivity assessment. Resistance to the pressure is considered 'High', and resilience 'High'. The sensitivity of this biotope to heavy deposition of up to 30cm of fine material added to the seabed in a single discreet event is assessed as 'Not Sensitive'.
Not assessed.
Electromagnetic changes
Underwater noise changes
Introduction of light or shading
Shading of the biotope (e.g. by construction of a pontoon, pier etc) could adversely affect the biotope in areas where the water clarity is also low, and tip the balance to shade tolerant species, resulting in the loss of the biotope directly within the shaded area, or a reduction in laminarian abundance from forest to park type biotopes.
Sensitivity assessment. Resistance is probably 'Low', with a 'Medium' resilience and a sensitivity of 'Medium', albeit with 'low' confidence due to the lack of direct evidence.
Barrier to species movement
Not relevant. this pressure is considered applicable to mobile species, e.g. fish and marine mammals rather than seabed habitats. Physical and hydrographic barriers may limit the dispersal of spores. But spore dispersal is not considered under the pressure definition and benchmark.
Death or injury by collision
Not relevant. Collision from grounding vessels is addressed under abrasion above.
Visual disturbance
Biological Pressures
Genetic modification & translocation of indigenous species
No evidence regarding the genetic modification or effects of translocation of native populations was found.
Introduction or spread of invasive non-indigenous species
Undaria pinnatifida has received a large amount of research attention as a major Invasive Non Indigenous Species (INIS) which could out-compete native UK kelp habitats (see Farrell & Fletcher, 2006; Thompson & Schiel, 2012, Brodie et al., 2014; Hieser et al., 2014). Undaria pinnatifida was first recorded in Plymouth Sound, UK in 2003 (NBN, 2015) subsequent surveys in 2011 have reported that Undaria pinnatifida is widespread throughout Plymouth Sound, colonizing rocky reef habitats. Where Undaria pinnatifida is present there was a significant decrease in the abundance of other Laminaria species, including Laminaria hyperborea (Heiser et al., 2014).
In New Zealand, Thompson & Schiel (2012) observed that native fucoids could out-compete U.pinnatifida and re-dominate the substratum. However, Thompson & Schiel (2012) suggested the fucoid recovery was partially due to an annual Undaria pinnatifida die back, which as noted by Heiser et al., (2014) did not occur in Plymouth Sound, UK. It is unknown whether Undaria pinnatifida will out-compete native macro-algae in the UK. However, from 2003-2011 Undaria pinnatifida had spread throughout Plymouth Sound, UK, becoming a visually dominant species at some locations within summer months (Hieser et al., 2014). While Undaria pinnatifida may replace Laminaria hyperborea in some locations within the UK, at the time of writing there is limited evidence available to assess what ecological impacts this invasion may have on Laminaria hyperborea associated communities e.g. red seaweeds.
Undaria pinnatifida was successfully eradicated on a sunken ship in Clatham Islands, New Zealand, by applying a heat treatment of 70 °C (see Wotton et al., 2004) however numerous other eradication attempts have failed, and as noted by Farrell & Fletcher (2006) once established Undaria pinadifida resists most attempts of long-term removal. The biotope is unlikely to fully recover until Undaria pinnatifida is fully removed from the habitat, which as stated above is unlikely to occur.
Sensitivity assessment. Resistance to the pressure is considered 'Low', and resilience 'Very Low'. The sensitivity of this biotope to introduction of INIS is assessed as 'High'.
Introduction of microbial pathogens
Galls on the blade of Laminaria hyperborea and spot disease are associated with the endophyte Streblonema sp. although the causal agent is unknown (bacteria, virus or endophyte). Resultant damage to the blade and stipe may increase losses in storms. The endophyte inhibits spore production and therefore recruitment and recoverability (Lein et al., 1991).
Sensitivity assessment. Resistance to the pressure is considered 'Medium', and resilience 'High'. The sensitivity of this biotope to introduction of microbial pathogens is assessed as 'Low'.
Removal of target species
Christie et al. (1998) observed Laminaria hyperborea habitat regeneration following commercial Laminaria hyperborea trawling in south Norway. Within the study area trawling removed all large canopy-forming adult L.hyperborea. Within 2-3 years of harvesting a new canopy had formed 1m off the seabed. The associated holdfast communities recovered in 6 years however the epiphytic stipe community did not fully recover within the same time period. Christie et al. (1998) suggested that kelp habitats were relatively resistant to direct disturbance of Laminaria hyperboreaa canopy.
Recurrent disturbance occurring at a smaller time scale than the recovery period of 2-6 years (stated above) could extend recovery time. Kain (1975) cleared sublittoral blocks of Laminaria hyperborea at different times of the year for several years. The first colonizers and succession community differed between blocks and at what time of year the blocks were cleared however within 2 years of clearance the blocks were dominated by Laminaria hyperborea. Leinaas & Christie (1996) also observed Laminaria hyperborea re-colonization of 'urchin barrens', following removal of urchins. The substratum was initially colinized by filamentous macroalgae and Saccharina latissima however after 2-4 years Laminaria hyperborea dominated the community.
Following disturbance or in areas were recurrent rapid disturbance occurs Laminaria hyperborea recruitment could also be affected by interspecifc competitive interactions with Non Indigenous Invasive Species or ephemeral algal species (Brodie et al., 2014; Smale et al., 2013), however evidence for this is limited and thus not included within this assessment.
Sensitivity assessment. Resistance to the pressure is considered 'None', and resilience 'Medium'. The sensitivity of this biotope to damage to seabed surface features is assessed as 'Medium'.
Removal of non-target species
Incidental/accidental removal of Laminaria hyperborea from other fisheries or extraction processes are likely to cause similar effects to that of direct harvesting; as such the same evidence has been used for both pressure assessments.
Christie et al., (1998) observed Laminaria hyperborea habitat regeneration following commercial Laminaria hyperborea trawling in south Norway. Within the study area trawling removed all large canopy-forming adult Laminaria hyperborea. Within 2-3 years of harvesting a new canopy had formed 1 m off the seabed. The associated holdfast communities recovered in six years however the epiphytic stipe community did not fully recover within the same time period. Christie et al. (1998) suggested that kelp habitats were relatively resistant to direct disturbance of Laminaria hyperborea canopy.
Recurrent disturbance occurring at a smaller time scale than the recovery period of 2-6 years (stated above) could extend recovery time. Kain (1975a) cleared sublittoral blocks of Laminaria hyperborea at different times of the year for several years. The first colonizers and succession community differed between blocks and at what time of year the blocks were cleared however within two years of clearance the blocks were dominated by Laminaria hyperborea. Leinaas & Christie (1996) also observed Laminaria hyperborea re-colonization of 'urchin barrens', following removal of urchins. The substratum was initially colinized by filamentous macroalgae and Saccharina latissima however after 2-4 years Laminaria hyperborea dominated the community.
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This review can be cited as:
Stamp, T.E. & Williams, E.. 2021. Laminaria hyperborea forest, foliose red seaweeds and a diverse fauna on tide-swept upper infralittoral rock. In Tyler-Walters H. Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 27-01-2023]. Available from: https://www.marlin.ac.uk/habitat/detail/227
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Sealife survey
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