information on the biology of species and the ecology of habitats found around the coasts and seas of the British Isles

Laminaria hyperborea with dense foliose red seaweeds on exposed infralittoral rock



UK and Ireland classification


Very exposed and exposed upper infralittoral bedrock or large boulders characterized by the kelp Laminaria hyperborea, beneath which is a dense turf of foliose red seaweeds. Three variations of this biotope have been described: the upper infralittoral kelp forest (EIR.LhypR.Ft), the kelp park below (EIR.LhypR.Pk) and a third type of kelp forest that is characterized by a mixture of Laminaria hyperborea and Laminaria ochroleuca (EIR.LhypR.Loch). The fauna of EIR.LhypR biotopes are markedly less abundant than kelp forests in areas of greater wave surge (EIR.LhypFa); sponges, anemones and polyclinid ascidians may be present, though never at high abundance. Beneath the understorey of red algae the rock surface is generally covered with encrusting coralline algae. (Information from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Depth range

0-5 m, 5-10 m, 10-20 m

Additional information


Listed By

Further information sources

Search on:

Habitat review


Ecological and functional relationships


Seasonal and longer term change


Habitat structure and complexity




Recruitment processes


Time for community to reach maturity


Additional information


Preferences & Distribution

Habitat preferences

Depth Range 0-5 m, 5-10 m, 10-20 m
Water clarity preferences
Limiting Nutrients Nitrogen (nitrates), Phosphorus (phosphates)
Salinity preferences Full (30-40 psu)
Physiographic preferences Open coast
Biological zone preferences Infralittoral
Substratum/habitat preferences Bedrock, Large to very large boulders, Small boulders
Tidal strength preferences Moderately Strong 1 to 3 knots (0.5-1.5 m/sec.), Weak < 1 knot (<0.5 m/sec.)
Wave exposure preferences Exposed, Extremely exposed, Very exposed
Other preferences

Additional Information

Van den Hoek (1982) suggested that the distribution of Laminaria hyperborea, and hence its associated biotope, was limited by temperatures between the 2 °C winter isotherm in the north and the 19 °C summer isotherm in the south.

Species composition

Species found especially in this biotope

Rare or scarce species associated with this biotope


Additional information

Little work on the rarity of species in kelp biotopes has been compiled (Birkett et al., 1998b). Kelp beds are diverse species rich habitats and over 1,800 species have been recorded in the UK kelp biotopes (Birkett et al., 1998b). Birkett et al. (1998b) list species recorded in UK biotope complexes by the MNCR (Appendix 5) together with common understorey algae and epiphytes (Appendices 4 & 3 respectively).
Holdfast fauna is a particularly species rich part of the biotope but no species have been suggested as specifically associated with holdfasts and therefore critical to the identity of the biotope.

Sensitivity review


Laminaria hyperborea provides substratum for numerous species and is the major source of primary production in this community, either directly or in the form of drift (broken off) algae. The biotope would cease to be EIR.LhypR if Laminaria hyperborea was lost. Helcion pellucidum grazes the blades of kelp directly and the laevis form excavates cavities in the holdfasts. These cavities weaken the holdfast making it more susceptible to removal by wave action and storms, and affects the age structure of the kelp population since older, larger plants are most likely to be lost. Sea-urchins have been responsible for removal of large areas of kelp bed in Nova Scotia, Norway and Southern California, resulting in 'urchin barrens'. The presence of this biotope is partly reliant on low or no populations of sea urchins, such as Echinus esculentus. Sea urchins are omnivores and graze drift algae, algal turfs, epifauna and epiphytes on kelp. Their grazing may prevent dominance by any one species, resulting in a more species rich epifauna/flora. Echinus esculentus has been shown to control the lower limit of Laminaria hyperborea in Isle of Man, affect the patchiness and species composition of the understorey algae and remove Laminaria hyperborea sporelings and juveniles. Sporelings did not survive past 2 years of age in areas grazed by urchins (Jones & Kain 1967; Kain 1979). Echinus esculentus is the common sea urchin in the British Isles. The EIR.LhypR biotope is characterized by dense red algae turf, of which Delesseria sanguinea is an important characterizing species. If the red algal turf was lost the biotope may change to EIR.LhypPar or EIR.LhypFa. Although sea urchins are considered keystone the mechanisms that control their populations are poorly understood. In undertaking this assessment of sensitivity, account is taken of knowledge of the biology of all characterizing species in the biotope. However, 'indicative species' are particularly important in undertaking the assessment because they have been subject to detailed research.

Species indicative of sensitivity

Community ImportanceSpecies nameCommon Name
Important characterizingDelesseria sanguineaSea beech
Key functionalEchinus esculentusEdible sea urchin
Key functionalHelcion pellucidumBlue-rayed limpet
Key structuralLaminaria hyperboreaTangle or cuvie

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High Moderate Moderate Decline Moderate
Key and characteristic species are highly intolerant of substratum loss. Removal of the bedrock or boulders will remove the kelp species and their gametophytes and cause more damage than harvesting alone. Intolerance has been assessed as high with a moderate recovery (see additional information).
Intermediate High Low Decline Low
Smothering by sediment e.g. 5 cm material for a month, is unlikely to damage Laminaria hyperborea plants but is likely to affect sporeling and gametophyte survival as well as holdfast fauna. A layer of sediment will interfere will 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 for 1 month. Once returned to normal conditions the gametophytes resumed growth or maturation within 1 month (very high recoverability) (tom Dieck, 1993). Intolerance to this factor is likely to be higher during the peak periods of sporulation or germling settlement. Understorey epifauna/flora may be adversely affected, especially suspension or filter feeding species and the settlement of larvae or spores may be impaired. Laminaria hyperborea, Delesseria sanguinea and Echinus esculentus were all assessed as of intermediate intolerance to smothering and accordingly, EIR.LhypR is assessed as of intermediate intolerance. Recoverability is likely to be high (see additional information).
Intermediate Very high Low Decline Low
Increased siltation may interfere with spore attachment, larval settlement and recruitment, smothering germlings and gametophytes (see above). Siltation may also reduce photosynthetic activity if deposited on lamina, and increase sediment scour of surfaces settled by algal spores or larvae (Fletcher, 1996). Fletcher (1996) reports that siltation in the vicinity of outfalls restricted the distribution of Laminaria spp. and resulted in the general absence or impoverishment of algae, leaving only a few selected species. It also increased the quantity of mussels which competed for space with benthic algae in the heavily polluted sites. Studies of cooling water discharge in a southern California Macrocystis forest suggested that turbidity and siltation significantly reduced the density of snails, sea urchins and starfish whereas two filter feeding species, a gorgonian coral and a sponge, increased in relative density (Birkett et al., 1998b). Exposed Laminaria hyperborea biotopes are likely to be free of silt and exhibit more foliose red algae than moderately exposed Laminaria hyperborea biotopes. Increased siltation may adversely affect recruitment in Patella pellucida and other grazing gastropods, and could result in decreases in population density resulting in increased abundance of epiphytes on stipes. Given the effect on settlement and recruitment, increased siltation may change the age structure of the algal population, reduce understorey flora/fauna species richness, and decrease gastropod grazing. Increased siltation may affect holdfast fauna, encouraging suspension feeders and silt tolerant communities (Moore, 1973a&b; Edwards, 1980). Sheppard et al. (1980) noted that increased suspended sediment (measured as clarity) reduced holdfast species diversity due to increased dominance of suspension feeders.
On balance, an intolerance of intermediate has been suggested, with very high recovery.
Tolerant Moderate Not relevant Minor decline Low
EIR.Lhyp.R is likely to be tolerant of a decrease in suspended sediment and it may exhibit a more diverse foliose red algae.
High Moderate Moderate Decline Moderate
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. Increased desiccation will probably remove more adult plants and their associated communities and red algae from the upper limit of its distribution. Most species associated with this biotope are intolerant of increased emergence and would be adversely affected. Although the majority of the kelp bed is subtidal and unlikely to be affected, EIR.LhypR has been assessed as being highly intolerant to desiccation to reflect the likely impact at the upper shore extent of this biotope. Recovery is expected to be moderate (see additional information).
High Moderate Moderate Decline Low
An increase in emergence of about 1 hour for a year will decrease the upper limit of the kelp beds, due to the increased insolation and risk of desiccation, with a concomitant decrease in species richness. The upper shore extent of the biotope may be replaced by sublittoral fringe biotopes, such as EIR.Ala and accordingly, intolerance has been assessed as high. Recovery is expected to be moderate (see additional information). The kelp park biotope (EIR.LhypR.Pk) is unlikely to be exposed to this factor due to its depth.
Tolerant* Not sensitive No change Low
A decrease in emergence may allow the kelp bed to extend its distribution further up the shore and tolerant* has been suggested.
High Moderate Moderate Minor decline Low
The morphology of the stipe and blade of kelps vary with water flow rate. Strong currents may lead to removal of kelps through detachment of holdfasts. Increased 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. Increased water flow rates may reduce the understorey epiflora, to be replaced by an epifauna dominated community (e.g. sponges, anemones and polclinid ascidians) as in the biotope EIR.LhypFa. The composition of the holdfast fauna may also change, e.g. energetic or sheltered water movements favour different species of amphipods (Moore, 1985). The recognisable biotope is likely to change significantly and, therefore, intolerance has been assessed as high with a moderate recovery (see additional information).
High Moderate Intermediate Decline Moderate
The morphology of the stipe and blade of kelps vary with water flow rate and the associated algal flora and suspension feeding faunal populations can also change significantly with different water flow regimes. However, EIR.LhypR is found in wave exposed and very wave exposed habitats and, therefore, wave energy is likely to far outweigh any reduction in water flow rate. The biotope may become more characteristic of moderately exposed habitats however overall, intolerance is likely to be low.
High Moderate Moderate Decline Low
The keystone species Laminaria hyperborea is stenothermal, with upper and lower lethal temperatures between 1-2 °C above or below the normal temperature tolerances of between 0 and 20 °C (depending on season) (Birkett et al., 1998b). Hoek (1982) suggests that Laminaria hyperborea can tolerate an annual temperature span of 17 °C with an upper and lower lethal temperatures of 19 °C and 2 °C. However, above 17 °C, gamete survival is reduced (Kain, 1971) and gametogenesis is inhibited at 21 °C in this species (tom Dieck, 1992). It is likely that the biotope as a whole will be damaged by temperatures outside the temperature tolerance of Laminaria hyperborea. Subtidal red algae are less tolerant of temperature extremes than intertidal Rhodophyceae, surviving between -2 °C (in seawater) and 18-23 °C (Lüning, 1990; Kain & Norton, 1990). Temperature increase may affect growth, recruitment or interfere with the reproductive cycle in some species. For example, there is some evidence to suggest that blade growth in Delesseria sanguinea is delayed until ambient sea temperatures fall below 13 °C, although blade growth is likely to be intrinsically linked to gametangia development (see Kain, 1987). Increases in temperature of e.g. 2 °C for a year or 5 °C for one week may raise ambient temperatures outside the tolerable range for the species within the biotope, causing changes in recruitment, growth rates and perhaps loss of red algae and changes in grazing patterns. It should be noted that increases in temperature tolerances by kelp species is less well tolerated in winter months than summer months (Birkett et al., 1998b).

Overall, EIR.LhypR has been assessed as being of high intolerance to increases in temperature at the benchmark level. Recovery is likely to be moderate (see additional information).

The sub-biotope EIR.LhypR.Loch is characterized by the presence of Laminaria ochroleuca, which is restricted to Devon, Cornwall and the Isles of Scilly but common on the coasts of Brittany. Long term increases of 1 -3 °C in temperature may allow Laminaria ochroleuca to spread northwards (Birkett et al., 1998b).
Low Very high Moderate No change Very low
The keystone species Laminaria hyperborea is stenothermal, with upper and lower lethal temperatures between 1-2 °C above or below the normal temperature tolerances of between 0 and 20 °C (depending on season) (Birkett et al., 1998b). Hoek (1982) suggests that Laminaria hyperborea can tolerate an annual temperature span of 17 °C with an upper and lower lethal temperatures of 19 °C and 2 °C. Given its distribution in the North Atlantic this species is likely to be tolerant of low temperatures. The temperature tolerances of the gametophyte stages are different to those of the adult. Gametophytic development has been observed at 0 °C although development is slow and suggests that 0 °C is close to the lowest temperature allowing vegetative development of the primary cells (Sjøtun & Schoschina, 2002). Subtidal red algae are less tolerant of temperature extremes than intertidal Rhodophyceae, surviving between -2 °C (in seawater) and 18-23 °C (Lüning, 1990; Kain & Norton, 1990). However, Delesseria sanguinea is a Northern species, likely to be tolerant of a decrease in temperature. Patella pellucida is also a Northern species, extending as far north as the northern Norway. Overseas, cold temperatures have been associated with high recruitment of sea urchins (Birkett et al., 1998b). Although the key structural, functional and characterizing species may be relatively tolerant of a decrease in temperature at the benchmark level, an intolerance of low has been suggested to reflect the likelihood that other may species commonly found in the biotope may be intolerant, in addition to the fact that the biotope is likely to be more intolerant to an acute drop in temperature than chronic change.
Intermediate Moderate Moderate Minor decline Moderate
Turbidity will primarily affect Laminaria hyperborea and affect the depth to which it is likely to grow. In the sub-biotope EIR.LhypR.Pk Laminaria hyperborea occurs at low density, due mainly to limiting light levels. Increased turbidity is likely to reduce the depth to which the kelp park extends, raising its upper limit and thereby decreasing the lower limit of the kelp forest. Red algae are shade tolerant extending to greater depths and probably not as intolerant of increases in turbidity as kelp species. Suspended material in the vicinity of outfalls has been reported to result in reduced depth range and fewer new plants under the canopy (Fletcher 1996) (see 'siltation' above). Intolerance has been assessed as intermediate. Recovery is likely to be moderate.
Tolerant* Not sensitive No change Moderate
A decrease in turbidity is likely to benefit the algal component of this biotope which may experience enhance primary productivity. Tolerant* has been suggested.
Intermediate Moderate Moderate Minor decline Moderate
Increased wave exposure is likely to remove older kelp plants, especially from the upper extent of the kelp forest, where Laminaria hyperborea may become replaced by kelps more tolerant of stronger wave action such as Laminaria digitata and Alaria esculenta. The extent of the biotope may be reduced and, therefore, intolerance has been assessed as intermediate.
High Moderate Intermediate Decline Moderate
Decreased wave exposure may not damage the Laminaria hyperborea forest or park, but may result in loss of foliose red algae and an increase in abundance of filamentous red algae, characteristic of the biotope MIR.Lhyp in which case the biotope will no longer be EIR.LhypR. In very wave sheltered situations Laminaria hyperborea is replaced by Saccharina latissima and the biotope may change significantly should the wave exposure drop from sheltered to very sheltered. Intolerance has been assessed as high with a moderate recovery (see additional information).
Tolerant Not relevant Not relevant Not relevant Very low
Fish species using the kelp beds as a nursery or feeding ground may be disturbed by underwater noise or vibration. However, little information on the effect of this factor on these species was available.
Tolerant Not relevant Not relevant Not relevant Very low
Fish species using the kelp beds as a nursery or feeding ground may be affected or disturbed by visual presence but little information was found.
Intermediate Moderate Moderate Decline Moderate
Laminarians and red algae are likely to be damaged abrasion due to anchor impact and sand or cobble scour. However, a passing scallop dredge is likely to remove or damage a proportion of the kelp and red algae present. Therefore, the community as a whole is likely to be of intermediate intolerance to physical disturbance at the benchmark level. . This biotope will be more intolerant of higher levels or frequency of physical disturbance e.g. routine or numerous anchorages. For recoverability, see additional information below.
High Moderate Moderate Decline Moderate
Laminaria hyperborea and most other algae cannot reattach once removed and will be lost. Cleared areas were colonized by opportunistic species such as Alaria esculenta, Saccorhiza polyschides, and Desmarestia spp., but were out-competed by Laminaria hyperborea within 3 years (Kain, 1975; Kain, 1979). Norwegian studies suggest that kelp communities take at least 10 years to recover from harvesting (Svendsen, 1972, cited in Birkett et al., 1998b). Similarly many species of epifauna have a permanent attachment and would be lost if displaced. However, species such as Echinus esculentus and Patella pellucida are relatively insensitive to displacement.

Chemical Pressures

Intermediate High Low Minor decline Moderate
Echinus esculentus was assessed as highly intolerant of synthetic chemicals. O'Brian & Dixon (1976) suggested that red algae were the most sensitive group of macrophytes to oil and dispersant contamination (see also 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 the 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 Spermothamnium repens and some tufts of Jania rubens survived together with Laurencia 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 loss of red algae would result in loss of characteristic species and the biotope would cease to be EIR.LhypR. Grazing gastropods and amphipods are likely to be intolerant of synthetic chemicals such as TBT and dispersants. This biotope is primarily subtidal and unlikely to be smothered by oil but its exposure to wave action may allow dispersant chemicals to penetrate deeper into the water column, as suggested by Smith (1968). However, surveys of subtidal communities at a number sites between 1- 22.5m below chart datum, including Laminaria hyperborea communities, showed no noticeable impacts of the Sea Empress oil spill and clean up (Rostron & Bunker, 1997). Therefore, although the kelp species themselves may be relatively unaffected by synthetic contaminants, characterizing red algae and Echinus esculentus, together with grazing gastropods may be highly intolerant, resulting in significant changes in species richness and community structure. However, red algae and urchins are likely to recover relatively quickly.
Heavy metal contamination
Intermediate High Low Minor decline Low
Bryan (1984) suggested that the general order for heavy metal toxicity in seaweeds is: Organic Hg > inorganic Hg > Cu > Ag > Zn > Cd > Pb. Cole et al. (1999) reported that Hg was very toxic to macrophytes. Similarly, Hopkin & Kain (1978) demonstrated sublethal affects of heavy metals of 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. Adult Echinus esculentus are long-lived and poor recruitment may not reduce grazing pressure in the short term. Although macrophyte 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.
Hydrocarbon contamination
Intermediate High Low Minor decline Moderate
The mucilaginous slime layer coating 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 forest. Similarly, surveys of subtidal communities at a number sites between 1 -22.5m below chart datum, including Laminaria hyperborea 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). Echinus esculentus populations in the vicinity of an oil terminal in A Coruna Bay, Spain showed developmental abnormalities in the skeleton. The tissues contained high levels of aliphatic hydrocarbons, naphthalenes, pesticides and heavy metals (Zn, Hg, Cd, Pb, and Cu) (Gomez & Miguez-Rodriguez, 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 and sea urchins 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. During recovery the community may appear similar to grazed kelp forest biotope MIR.LhypGz.
Radionuclide contamination
No information No information No information Insufficient
Not relevant
Changes in nutrient levels
Intermediate Moderate Moderate Decline Low
Holt et al. (1995) suggest that Laminaria hyperborea may be tolerant of eutrophication since healthy populations are found at ends of sublittoral untreated sewage outfalls in the Isle of Man. Nutrients may be added to macrophyte cultures to increase productivity. 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 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 (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.
Not relevant Not relevant Not relevant Not relevant Not relevant
The full salinity, infralittoral habitat within which this biotope occurs is unlikely to experience significant increases in salinity and this factor is considered to be irrelevant.
Intermediate Moderate Intermediate Minor decline Low
Lüning (1990) suggest that kelps are stenohaline, their tolerance to salinity covering 16 - 50 psu over a 24 hr period. Optimal growth probably occurs between 30 -35 psu and growth rates are likely to be affected by periodic salinity stress. Hopkin & Kain (1978) stated that Laminaria hyperborea early sporophytes grew optimally between 20 -35 psu but did not survive at 6 psu. The representative species suggested for this biotope are assessed as intermediate intolerance to reduced salinity. Birkett et al. (1998) suggest that long term changes in salinity may result in loss of affected kelp beds and, therefore, loss of this biotope. Accordingly, intolerance has been assessed as intermediate with a moderate recoverability (see additional information).
Intermediate High Low Decline Very low
The effects of deoxygenation in plants has been little studied. Since plants produce oxygen they may be considered relatively insensitive. However, they may be more intolerant during darkness when they continue to respire. A study of the effects of anaerobiosis (no oxygen) on some marine algae concluded that Delesseria sanguinea was very intolerant of anaerobic conditions; at 15 °C death occurs within 24hrs and no recovery takes place although specimens survived at 5 °C (Hammer, 1972). Low concentrations of dissolved oxygen may be detrimental, especially to sedentary benthic epifauna and some species of red algae. A reduction in dissolved oxygen levels to 2mg/l will probably adversely affect several members of the epifauna and holdfast fauna, although this may be a rare occurrence in exposed conditions.

Biological Pressures

Intermediate High Low Minor decline Moderate
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. Echinus esculentus is susceptible to 'Bald-sea-urchin disease', which causes lesions, loss of spines, tube feet, pedicellariae, destruction of the upper layer of skeletal tissue and death. Bald sea-urchin disease was recorded from Echinus esculentus on the Brittany Coast and, although associated with mass mortalities of Strongylocentrotus franciscanus in California and Paracentrotus lividus in the French Mediterranean (Bower, 1996), no evidence of mass moralities of Echinus esculentus associated with disease have been recorded in Britain and Ireland.
Intermediate High Low Minor decline Very low
The Japanese kelp Undaria pinnatifida(wakame) has recently spread to south coast of England from northern Brittany and it thought to compete with native Saccorhiza polyschides. Macrocystis pyrifera was briefly introduced to French waters, for aquaculture, before it was stopped by international pressure. Macrocystis pyrifera is large and rapid growing and could potentially compete with native kelp species (Birkett et al., 1998b) resulting in different biotopes.
Intermediate Moderate Moderate Decline Moderate
Both Laminaria hyperborea and Echinus esculentus may be extracted. Removal of urchin predators such as lobsters or crawfish has been implicated in increases in urchin populations and therefore 'urchin barrens' and the loss of kelp beds. Similarly removal of grazing abalone by fishing is thought to have resulted in the loss of kelp beds as sea urchins populations benefited from reduced competition for food. However, attempts to correlate sea urchin numbers with specific predators are equivocal (Elner & Vadas, 1990; Birkett et al., 1998; Raffaelli & Hawkins, 1999). It is likely that there is a complex interaction between sea urchin numbers, recruitment and predation. Populations of Echinus esculentus, for example, are probably controlled by several predators, parasites, disease and recruitment. However, removal of predators or other grazers may perturb the community, making it more intolerant of natural fluctuations in sea urchin numbers or other perturbations and may result in loss of areas of kelp and 'urchin barrens'. However, extraction of Echinus esculentus may encourage dominant macroalgae, including kelps, and reduce the species richness of epiphyte and understorey fauna and flora.

Overall, an intolerance of intermediate has been suggested with a moderate recovery. Research on harvested populations of Laminaria hyperborea in Norway suggest that epiphytic and understorey fauna and flora were reduced in harvested areas compared to areas 10 years post harvesting (Sivertsen, 1991and Rinde et al., 1992, cited in Birkett et al., 1998b). Likewise recovery of the community may take at least 10 years (see 'General biology' page).

High Moderate Moderate Decline Moderate

Additional information

Although the kelp plants themselves may recover in 3-4 years, the community as a whole will take at least 10 years to return (Kain, 1979; Birkett et al., 1998b). Therefore, recoverability of the biotope from factors to which it has a high intolerance (and, in some cases, intermediate intolerance), will be moderate.

Wave exposed kelp forests with dense foliose seaweeds are likely to be more intolerant of incremental grazing presence and to reduction in wave exposure.


  1. Beszczynska-Möller, A., & Dye, S.R., 2013. ICES Report on Ocean Climate 2012. In ICES Cooperative Research Report, vol. 321 pp. 73.

  2. Birkett, D.A., Maggs, C.A., Dring, M.J. & Boaden, P.J.S., 1998b. Infralittoral reef biotopes with kelp species: an overview of dynamic and sensitivity characteristics for conservation management of marine SACs. Natura 2000 report prepared by Scottish Association of Marine Science (SAMS) for the UK Marine SACs Project., Scottish Association for Marine Science. (UK Marine SACs Project, vol V.). Available from:

  3. Bishop, G.M., 1985. Aspects of the reproductive ecology of the sea urchin Echinus esculentus L. Ph.D. thesis, University of Exeter, UK.

  4. Boney, A.D., 1971. Sub-lethal effects of mercury on marine algae. Marine Pollution Bulletin, 2, 69-71.

  5. Bower, S.M., 1996. Synopsis of Infectious Diseases and Parasites of Commercially Exploited Shellfish: Bald-sea-urchin Disease. [On-line]. Fisheries and Oceans Canada. [cited 26/01/16]. Available from:

  6. Brodie J., Williamson, C.J., Smale, D.A., Kamenos, N.A., Mieszkowska, N., Santos, R., Cunliffe, M., Steinke, M., Yesson, C. & Anderson, K.M., 2014. The future of the northeast Atlantic benthic flora in a high CO2 world. Ecology and Evolution, 4 (13), 2787-2798.

  7. Bryan, G.W., 1984. Pollution due to heavy metals and their compounds. In Marine Ecology: A Comprehensive, Integrated Treatise on Life in the Oceans and Coastal Waters, vol. 5. Ocean Management, part 3, (ed. O. Kinne), pp.1289-1431. New York: John Wiley & Sons.

  8. Burrows, M.T., Smale, D., O’Connor, N., Rein, H.V. & Moore, P., 2014. Marine Strategy Framework Directive Indicators for UK Kelp Habitats Part 1: Developing proposals for potential indicators. Joint Nature Conservation Comittee,  Peterborough. Report no. 525.

  9. Casas, G., Scrosati, R. & Piriz, M.L., 2004. The invasive kelp Undaria pinnatifida (Phaeophyceae, Laminariales) reduces native seaweed diversity in Nuevo Gulf (Patagonia, Argentina). Biological Invasions, 6 (4), 411-416.

  10. Christie, H., Fredriksen, S. & Rinde, E., 1998. Regrowth of kelp and colonization of epiphyte and fauna community after kelp trawling at the coast of Norway. Hydrobiologia, 375/376, 49-58.

  11. Cole, S., Codling, I.D., Parr, W., Zabel, T., 1999. Guidelines for managing water quality impacts within UK European marine sites [On-line]. UK Marine SACs Project. [Cited 26/01/16]. Available from:

  12. Connell, S.D., 2003. The monopolization of understorey habitat by subtidal encrusting coralline algae: a test of the combined effects of canopy-mediated light and sedimentation. Marine Biology, 142 (6), 1065-1071.

  13. Connor, D.W., Allen, J.H., Golding, N., Howell, K.L., Lieberknecht, L.M., Northen, K.O. & Reker, J.B., 2004. The Marine Habitat Classification for Britain and Ireland. Version 04.05. Joint Nature Conservation Committee, Peterborough.

  14. Connor, D.W., Dalkin, M.J., Hill, T.O., Holt, R.H.F. & Sanderson, W.G., 1997a. Marine biotope classification for Britain and Ireland. Vol. 2. Sublittoral biotopes. Joint Nature Conservation Committee, Peterborough, JNCC Report no. 230, Version 97.06., Joint Nature Conservation Committee, Peterborough, JNCC Report no. 230, Version 97.06.

  15. Crisp, D.J. (ed.), 1964. The effects of the severe winter of 1962-63 on marine life in Britain. Journal of Animal Ecology, 33, 165-210.

  16. Dauvin, J.C., Bellan, G., Bellan-Santini, D., Castric, A., Francour, P., Gentil, F., Girard, A., Gofas, S., Mahe, C., Noel, P., & Reviers, B. de., 1994. Typologie des ZNIEFF-Mer. Liste des parametres et des biocoenoses des cotes francaises metropolitaines. 2nd ed. Secretariat Faune-Flore, Museum National d'Histoire Naturelle, Paris (Collection Patrimoines Naturels, Serie Patrimoine Ecologique, No. 12). Coll. Patrimoines Naturels, vol. 12, Secretariat Faune-Flore, Paris.

  17. Davies, C.E. & Moss, D., 1998. European Union Nature Information System (EUNIS) Habitat Classification. Report to European Topic Centre on Nature Conservation from the Institute of Terrestrial Ecology, Monks Wood, Cambridgeshire. [Final draft with further revisions to marine habitats.], Brussels: European Environment Agency.

  18. Dayton, P.K., Tegner, M.J., Parnell, P.E. & Edwards, P.B., 1992. Temporal and spatial patterns of disturbance and recovery in a kelp forest community. Ecological Monographs, 62, 421-445.

  19. Devlin, M.J., Barry, J., Mills, D.K., Gowen, R.J., Foden, J., Sivyer, D. & Tett, P., 2008. Relationships between suspended particulate material, light attenuation and Secchi depth in UK marine waters. Estuarine, Coastal and Shelf Science, 79 (3), 429-439.

  20. Dieck, T.I., 1992. North Pacific and North Atlantic digitate Laminaria species (Phaeophyta): hybridization experiments and temperature responses. Phycologia, 31, 147-163.

  21. Dieck, T.I., 1993. Temperature tolerance and survival in darkness of kelp gametophytes (Laminariales: Phaeophyta) - ecological and biogeographical implications. Marine Ecology Progress Series, 100, 253-264.

  22. Dinnel, P.A., Pagano, G.G., & Oshido, P.S., 1988. A sea urchin test system for marine environmental monitoring. In Echinoderm Biology. Proceedings of the Sixth International Echinoderm Conference, Victoria, 23-28 August 1987, (R.D. Burke, P.V. Mladenov, P. Lambert, Parsley, R.L. ed.), pp 611-619. Rotterdam: A.A. Balkema.

  23. Eckman, J.E., Duggins, D.O., Sewell, A.T., 1989. Ecology of understory kelp environments. I. Effects of kelps on flow and particle transport near the bottom. Journal of Experimental Marine Biology and Ecology, 129, 173-187.

  24. Edwards, A., 1980. Ecological studies of the kelp Laminaria hyperborea and its associated fauna in south-west Ireland. Ophelia, 9, 47-60.

  25. Elner, R.W. & Vadas, R.L., 1990. Inference in ecology: the sea urchin phenomenon in the northwest Atlantic. American Naturalist, 136, 108-125.

  26. Erwin, D.G., Picton, B.E., Connor, D.W., Howson, C.M., Gilleece, P. & Bogues, M.J., 1990. Inshore Marine Life of Northern Ireland. Report of a survey carried out by the diving team of the Botany and Zoology Department of the Ulster Museum in fulfilment of a contract with Conservation Branch of the Department of the Environment (N.I.)., Ulster Museum, Belfast: HMSO.

  27. Farrell, P. & Fletcher, R., 2006. An investigation of dispersal of the introduced brown alga Undaria pinnatifida (Harvey) Suringar and its competition with some species on the man-made structures of Torquay Marina (Devon, UK). Journal of Experimental Marine Biology and Ecology, 334 (2), 236-243.

  28. Fletcher, R.L., 1996. The occurrence of 'green tides' - a review. In Marine Benthic Vegetation. Recent changes and the Effects of Eutrophication (ed. W. Schramm & P.H. Nienhuis). Berlin Heidelberg: Springer-Verlag. [Ecological Studies, vol. 123].

  29. Fredriksen, S., Sjøtun, K., Lein, T.E. & Rueness, J., 1995. Spore dispersal in Laminaria hyperborea (Laminariales, Phaeophyceae). Sarsia, 80 (1), 47-53.

  30. Frieder, C., Nam, S., Martz, T. & Levin, L., 2012. High temporal and spatial variability of dissolved oxygen and pH in a nearshore California kelp forest. Biogeosciences, 9 (10), 3917-3930.

  31. Gommez, J.L.C. & Miguez-Rodriguez, L.J., 1999. Effects of oil pollution on skeleton and tissues of Echinus esculentus L. 1758 (Echinodermata, Echinoidea) in a population of A Coruna Bay, Galicia, Spain. In Echinoderm Research 1998. Proceedings of the Fifth European Conference on Echinoderms, Milan, 7-12 September 1998, (ed. M.D.C. Carnevali & F. Bonasoro) pp. 439-447. Rotterdam: A.A. Balkema.

  32. Gorman, D., Bajjouk, T., Populus, J., Vasquez, M. & Ehrhold, A., 2013. Modeling kelp forest distribution and biomass along temperate rocky coastlines. Marine Biology, 160 (2), 309-325.

  33. Grandy, N., 1984. The effects of oil and dispersants on subtidal red algae. Ph.D. Thesis. University of Liverpool.

  34. Hammer, L., 1972. Anaerobiosis in marine algae and marine phanerograms. In Proceedings of the Seventh International Seaweed Symposium, Sapporo, Japan, August 8-12, 1971 (ed. K. Nisizawa, S. Arasaki, Chihara, M., Hirose, H., Nakamura V., Tsuchiya, Y.), pp. 414-419. Tokyo: Tokyo University Press.

  35. Harkin, E., 1981. Fluctuations in epiphyte biomass following Laminaria hyperborea canopy removal. In Proceedings of the Xth International Seaweed Symposium, Gø teborg, 11-15 August 1980 (ed. T. Levring), pp.303-308. Berlin: Walter de Gruyter.

  36. Hayward, P.J. 1988. Animals on seaweed. Richmond, Surrey: Richmond Publishing Co. Ltd. [Naturalists Handbooks 9].

  37. Heiser, S., Hall-Spencer, J.M. & Hiscock, K., 2014. Assessing the extent of establishment of Undaria pinnatifida in Plymouth Sound Special Area of Conservation, UK. Marine Biodiversity Records, 7, e93.

  38. Hiscock, K. & Mitchell, R., 1980. The Description and Classification of Sublittoral Epibenthic Ecosystems. In The Shore Environment, Vol. 2, Ecosystems, (ed. J.H. Price, D.E.G. Irvine, & W.F. Farnham), 323-370. London and New York: Academic Press. [Systematics Association Special Volume no. 17(b)].

  39. Hoare, R. & Hiscock, K., 1974. An ecological survey of the rocky coast adjacent to the effluent of a bromine extraction plant. Estuarine and Coastal Marine Science, 2 (4), 329-348.

  40. Holt, T.J., Jones, D.R., Hawkins, S.J. & Hartnoll, R.G., 1995. The sensitivity of marine communities to man induced change - a scoping report. Countryside Council for Wales, Bangor, Contract Science Report, no. 65.

  41. Hopkin, R. & Kain, J.M., 1978. The effects of some pollutants on the survival, growth and respiration of Laminaria hyperborea. Estuarine and Coastal Marine Science, 7, 531-553.

  42. JNCC, 2015. The Marine Habitat Classification for Britain and Ireland Version 15.03. JNCC: JNCC. 2015(20/05/2015).

  43. JNCC (Joint Nature Conservation Committee), 1999. Marine Environment Resource Mapping And Information Database (MERMAID): Marine Nature Conservation Review Survey Database. [on-line]

  44. Johansson ,G., Eriksson, B.K., Pedersen, M. & Snoeijs, P., 1998. Long term changes of macroalgal vegetation in the Skagerrak area. Hydrobiologia, 385, 121-138.

  45. Jones, C.G., Lawton, J.H. & Shackak, M., 1994. Organisms as ecosystem engineers. Oikos, 69, 373-386.

  46. Jones, D.J., 1971. Ecological studies on macro-invertebrate communities associated with polluted kelp forest in the North Sea. Helgolander Wissenschaftliche Meersuntersuchungen, 22, 417-431.

  47. Jones, L.A., Hiscock, K. & Connor, D.W., 2000. Marine habitat reviews. A summary of ecological requirements and sensitivity characteristics for the conservation and management of marine SACs. Joint Nature Conservation Committee, Peterborough. (UK Marine SACs Project report.). Available from:

  48. Jones, N.S. & Kain, J.M., 1967. Subtidal algal recolonisation following removal of Echinus. Helgolander Wissenschaftliche Meeresuntersuchungen, 15, 460-466.

  49. Kain, J.M., 1964. Aspects of the biology of Laminaria hyperborea III. Survival and growth of gametophytes. Journal of the Marine Biological Association of the United Kingdom, 44 (2), 415-433.

  50. Kain, J.M. & Svendsen, P., 1969. A note on the behaviour of Patina pellucida in Britain and Norway. Sarsia, 38, 25-30.

  51. Kain, J.M., 1971a. Synopsis of biological data on Laminaria hyperborea. FAO Fisheries Synopsis, no. 87.

  52. Kain, J.M., 1975a. Algal recolonization of some cleared subtidal areas. Journal of Ecology, 63, 739-765.

  53. Kain, J.M., 1979. A view of the genus Laminaria. Oceanography and Marine Biology: an Annual Review, 17, 101-161.

  54. Kain, J.M., 1987. Photoperiod and temperature as triggers in the seasonality of Delesseria sanguinea. Helgolander Meeresuntersuchungen, 41, 355-370.

  55. Kain, J.M., & Norton, T.A., 1990. Marine Ecology. In Biology of the Red Algae, (ed. K.M. Cole & Sheath, R.G.). Cambridge: Cambridge University Press.

  56. Kain, J.M., Drew, E.A. & Jupp, B.P., 1975. Light and the ecology of Laminaria hyperborea II. In Proceedings of the Sixteenth Symposium of the British Ecological Society, 26-28 March 1974. Light as an Ecological Factor: II (ed. G.C. Evans, R. Bainbridge & O. Rackham), pp. 63-92. Oxford: Blackwell Scientific Publications.

  57. Karsten, U., 2007. Research note: salinity tolerance of Arctic kelps from Spitsbergen. Phycological Research, 55 (4), 257-262.

  58. Kinne, O. (ed.), 1984. Marine Ecology: A Comprehensive, Integrated Treatise on Life in Oceans and Coastal Waters.Vol. V. Ocean Management Part 3: Pollution and Protection of the Seas - Radioactive Materials, Heavy Metals and Oil. Chichester: John Wiley & Sons.

  59. Kinne, O., 1977. International Helgoland Symposium "Ecosystem research": summary, conclusions and closing. Helgoländer Wissenschaftliche Meeresuntersuchungen, 30(1-4), 709-727.

  60. Kitching, J., 1941. Studies in sublittoral ecology III. Laminaria forest on the west coast of Scotland; a study of zonation in relation to wave action and illumination. The Biological Bulletin, 80 (3), 324-337

  61. Kregting, L., Blight, A., Elsäßer, B. & Savidge, G., 2013. The influence of water motion on the growth rate of the kelp Laminaria hyperborea. Journal of Experimental Marine Biology and Ecology, 448, 337-345.

  62. Krumhansl, K.A., 2012. Detrital production in kelp beds. degree of Doctor of Philosophy, Department of Biology, Dalhousie University, Halifax, Nova Scotia.

  63. Kruuk, H., Wansink, D. & Moorhouse, A., 1990. Feeding patches and diving success of otters, Lutra lutra, in Shetland. Oikos, 57, 68-72.

  64. Lang, C. & Mann, K., 1976. Changes in sea urchin populations after the destruction of kelp beds. Marine Biology, 36 (4), 321-326.

  65. Lein, T.E, Sjotun, K. & Wakili, S., 1991. Mass - occurrence of a brown filamentous endophyte in the lamina of the kelp Laminaria hyperborea (Gunnerus) Foslie along the south western coast of Norway Sarsia, 76, 187-193.

  66. Leinaas, H.P. & Christie, H., 1996. Effects of removing sea urchins (Strongylocentrotus droebachiensis): stability of the barren state and succession of kelp forest recovery in the east Atlantic. Oecologia, 105(4), 524-536.

  67. Lobban, C.S. & Harrison, P.J., 1997. Seaweed ecology and physiology. Cambridge: Cambridge University Press.

  68. Lüning, K., 1990. Seaweeds: their environment, biogeography, and ecophysiology: John Wiley & Sons.

  69. Mann, K.H., 1982. Kelp, sea urchins, and predators: a review of strong interactions in rocky subtidal systems of eastern Canada, 1970-1980. Netherlands Journal of Sea Research, 16, 414-423.

  70. Miller III, H.L., Neale, P.J. & Dunton, K.H., 2009. Biological weighting functions for UV inhibtion of photosynthesis in the kelp Laminaria hyperborea (Phaeophyceae) 1. Journal of Phycology, 45 (3), 571-584.

  71. Moore, P.G., 1973a. The kelp fauna of north east Britain I. Function of the physical environment. Journal of Experimental Marine Biology and Ecology, 13, 97-125.

  72. Moore, P.G., 1973b. The kelp fauna of north east Britain. II. Multivariate classification: turbidity as an ecological factor. Journal of Experimental Marine Biology and Ecology, 13, 127-163.

  73. Moore, P.G., 1978. Turbidity and kelp holdfast Amphipoda. I. Wales and S.W. England. Journal of Experimental Marine Biology and Ecology, 32, 53-96.

  74. Moore, P.G., 1985. Levels of heterogeneity and the amphipod fauna of kelp holdfasts. In The Ecology of Rocky Coasts: essays presented to J.R. Lewis, D.Sc. (ed. P.G. Moore & R. Seed), 274-289. London: Hodder & Stoughton Ltd.

  75. NBN, 2015. National Biodiversity Network 2015(20/05/2015).

  76. Nichols, D., 1981. The Cornish Sea-urchin Fishery. Cornish Studies, 9, 5-18.

  77. Norderhaug, K., 2004. Use of red algae as hosts by kelp-associated amphipods. Marine Biology, 144 (2), 225-230.

  78. Norderhaug, K.M. & Christie, H., 2011. Secondary production in a Laminaria hyperborea kelp forest and variation according to wave exposure. Estuarine, Coastal and Shelf Science, 95 (1), 135-144.

  79. Norderhaug, K.M. & Christie, H.C., 2009. Sea urchin grazing and kelp re-vegetation in the NE Atlantic. Marine Biology Research, 5 (6), 515-528.

  80. Norderhaug, K.M., Christie, H. & Fredriksen, S., 2007. Is habitat size an important factor for faunal abundances on kelp (Laminaria hyperborea)? Journal of Sea Research, 58 (2), 120-124.

  81. Nordheim, van, H., Andersen, O.N. & Thissen, J., 1996. Red lists of Biotopes, Flora and Fauna of the Trilateral Wadden Sea area, 1995. Helgolander Meeresuntersuchungen, 50 (Suppl.), 1-136.

  82. Norton, T.A., 1978. The factors influencing the distribution of Saccorhiza polyschides in the region of Lough Ine. Journal of the Marine Biological Association of the United Kingdom, 58, 527-536.

  83. Norton, T.A., 1992. Dispersal by macroalgae. British Phycological Journal, 27, 293-301.

  84. Norton, T.A., Hiscock, K. & Kitching, J.A., 1977. The Ecology of Lough Ine XX. The Laminaria forest at Carrigathorna. Journal of Ecology, 65, 919-941.

  85. O'Brien, P.J. & Dixon, P.S., 1976. Effects of oils and oil components on algae: a review. British Phycological Journal, 11, 115-142.

  86. Pedersen, M.F., Nejrup, L.B., Fredriksen, S., Christie, H. & Norderhaug, K.M., 2012. Effects of wave exposure on population structure, demography, biomass and productivity of the kelp Laminaria hyperborea. Marine Ecology Progress Series, 451, 45-60.

  87. Penfold, R., Hughson, S., & Boyle, N., 1996. The potential for a sea urchin fishery in Shetland., 2000-04-14

  88. Philippart, C.J., Anadón, R., Danovaro, R., Dippner, J.W., Drinkwater, K.F., Hawkins, S.J., Oguz, T., O'Sullivan, G. & Reid, P.C., 2011. Impacts of climate change on European marine ecosystems: observations, expectations and indicators. Journal of Experimental Marine Biology and Ecology, 400 (1), 52-69.

  89. Raffaelli, D. & Hawkins, S., 1999. Intertidal Ecology 2nd edn.. London: Kluwer Academic Publishers.

  90. Rietema, H., 1993. Ecotypic differences between Baltic and North Sea populations of Delesseria sanguinea and Membranoptera alata. Botanica Marina, 36, 15-21.

  91. Rinde, E. & Sjøtun, K., 2005. Demographic variation in the kelp Laminaria hyperborea along a latitudinal gradient. Marine Biology, 146 (6), 1051-1062.

  92. Rinde, E., Christie, H., Fredriksen, S. & Sivertsen, A., 1992. Ecological consequences of kelp trawling: Importance of the structure of the kelp forest for abundance of fauna in the kelp holdfasts, benthic fauna and epiphytes. Norsk Institutt for Naturforskning. Oppdragsmelding, (127), 1-37.

  93. Rostron, D.M. & Bunker, F. St P.D., 1997. An assessment of sublittoral epibenthic communities and species following the Sea Empress oil spill. A report to the Countryside Council for Wales from Marine Seen & Sub-Sea Survey., Countryside Council for Wales, Bangor, CCW Sea Empress Contact Science, no. 177.

  94. Schiel, D.R. & Foster, M.S., 1986. The structure of subtidal algal stands in temperate waters. Oceanography and Marine Biology: an Annual Review, 24, 265-307.

  95. Sheppard, C.R.C. & Bellamy, D.J., 1974. Pollution of the Mediterranean around Naples. Marine Pollution Bulletin, 5, 42-44.

  96. Sheppard, C.R.C., Bellamy, D.J. & Sheppard, A.L.S., 1980. Study of the fauna inhabiting the holdfasts of Laminaria hyperborea (Gunn.) Fosl. along some environmental and geographical gradients. Marine Environmental Research, 4, 25-51.

  97. Sivertsen, K., 1997. Geographic and environmental factors affecting the distribution of kelp beds and barren grounds and changes in biota associated with kelp reduction at sites along the Norwegian coast. Canadian Journal of Fisheries and Aquatic Sciences, 54, 2872-2887.

  98. Sjøtun, K., Christie, H. & Helge Fosså, J., 2006. The combined effect of canopy shading and sea urchin grazing on recruitment in kelp forest (Laminaria hyperborea). Marine Biology Research, 2 (1), 24-32.

  99. Sjøtun, K. & Fredriksen, S., 1995. Growth allocation in Laminaria hyperborea (Laminariales, Phaeophyceae) in relation to age and wave exposure. Marine Ecology Progress Series, 126, 213-222.

  100. Sjøtun, K. & Schoschina, E.V., 2002. Gametophytic development of Laminaria spp. (Laminariales, Phaeophyta) at low temperatures. Phycologia, 41, 147-152.

  101. Sjøtun, K., Fredriksen, S. & Rueness, J., 1998. Effect of canopy biomass and wave exposure on growth in Laminaria hyperborea (Laminariaceae: Phaeophyta). European Journal of Phycology, 33, 337-343.

  102. Smale, D.A., Burrows, M.T., Moore, P., O'Connor, N. & Hawkins, S.J., 2013. Threats and knowledge gaps for ecosystem services provided by kelp forests: a northeast Atlantic perspective. Ecology and evolution, 3 (11), 4016-4038.

  103. Smale, D.A., Wernberg, T., Yunnie, A.L. & Vance, T., 2014. The rise of Laminaria ochroleuca in the Western English Channel (UK) and comparisons with its competitor and assemblage dominant Laminaria hyperborea. Marine ecology.

  104. Smith, J.E. (ed.), 1968. 'Torrey Canyon'. Pollution and marine life. Cambridge: Cambridge University Press.

  105. Somerfield, P.J. & Warwick, R.M., 1999. Appraisal of environmental impact and recovery using Laminaria holdfast faunas. Sea Empress, Environmental Evaluation Committee., Countryside Council for Wales, Bangor, CCW Sea Empress Contract Science, Report no. 321.

  106. Staehr, P.A. & Wernberg, T., 2009. Physiological responses of Ecklonia radiata (Laminariales) to a latitudinal gradient in ocean temperature. Journal of Phycology, 45, 91-99.

  107. Steneck, R.S., Graham, M.H., Bourque, B.J., Corbett, D., Erlandson, J.M., Estes, J.A. & Tegner, M.J., 2002. Kelp forest ecosystems: biodiversity, stability, resilience and future. Environmental conservation, 29 (04), 436-459.

  108. Steneck, R.S., Vavrinec, J. & Leland, A.V., 2004. Accelerating trophic-level dysfunction in kelp forest ecosystems of the western North Atlantic. Ecosystems, 7 (4), 323-332.

  109. Stock, J.H., 1988. Lamippidae (Copepoda : Siphonostomatoida) parasitic in Alcyonium. Journal of the Marine Biological Association of the United Kingdom, 68, 351-359.

  110. Thompson, G.A. & Schiel, D.R., 2012. Resistance and facilitation by native algal communities in the invasion success of Undaria pinnatifida. Marine Ecology, Progress Series, 468, 95-105.

  111. Vadas, R.L. & Elner, R.W., 1992. Plant-animal interactions in the north-west Atlantic. In Plant-animal interactions in the marine benthos, (ed. D.M. John, S.J. Hawkins & J.H. Price), 33-60. Oxford: Clarendon Press. [Systematics Association Special Volume, no. 46].

  112. Vadas, R.L., Johnson, S. & Norton, T.A., 1992. Recruitment and mortality of early post-settlement stages of benthic algae. British Phycological Journal, 27, 331-351.

  113. Van den Hoek, C., 1982. The distribution of benthic marine algae in relation to the temperature regulation of their life histories. Biological Journal of the Linnean Society, 18, 81-144.

  114. Vost, L.M., 1983. The influence of Echinus esculentus grazing on subtidal algal communities. British Phycological Journal, 18, 211.

  115. Walker, F. T., 1953. Distribution of Laminariaceae around Scotland. Journal du Conseil, 20 (2), 160-166.

  116. Whittick, A., 1983. Spatial and temporal distributions of dominant epiphytes on the stipes of Laminaria hyperborea (Gunn.) Fosl. (Phaeophyta: Laminariales) in S.E. Scotland. Journal of Experimental Marine Biology and Ecology, 73, 1-10.

  117. Wilkinson, M., 1995. Information review on the impact of kelp harvesting. Scottish Natural Heritage Review, no. 34, 54 pp.

  118. Wotton, D.M., O'Brien, C., Stuart, M.D. & Fergus, D.J., 2004. Eradication success down under: heat treatment of a sunken trawler to kill the invasive seaweed Undaria pinnatifida. Marine Pollution Bulletin, 49 (9), 844-849.


This review can be cited as:

Stamp, T.E. & Tyler-Walters, H. 2015. [Laminaria hyperborea] with dense foliose red seaweeds on exposed infralittoral rock. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. [cited 19-06-2018]. Available from:

Last Updated: 30/11/2015