|Researched by||Dr Harvey Tyler-Walters||Refereed by||Dr Joanna Jones|
|Authority||(Gunnerus) Foslie, 1884|
|Other common names||-||Synonyms||-|
A large conspicuous kelp which can grow up to 3.5 m in length in suitable conditions although this length is rarely attained (J. Jones, pers. comm.). The blade is broad, large, tough, flat and divided into 5 - 20 straps or fingers (digitate). The blade is glossy, golden brown to very dark brown in colour. The holdfast is large, conical and branched with conspicuous haptera. The stipe is stiff, rough textured, thick at the base and tapers towards the frond. The stipe stands erect when out of water. The stipe is often covered with numerous epifauna and epiflora. The amount of energy allocated to growth of the stipe, and consequently maximum length of stipe, varies with season, the age of plant and location. This species is often confused with Laminaria digitata, especially when young.
Other common names include, redware, cuvy, sea rod, mayweed or Slat mara. The new blade grows below the older from November onwards. The old blade is shed in spring and early summer. Blade and stipe vary with exposure and current. In sheltered conditions the blade has few or no digits and the stipe becomes thin but in exposed conditions the blade is deeply digitate and the stipe becomes thick. The stipe is usually up to 1m long but stipes up to 3m long have been recorded (Parke unpublished, cited in Kain, 1971a).
- none -
|Phylum||Ochrophyta||Brown and yellow-green seaweeds|
|Authority||(Gunnerus) Foslie, 1884|
|Typical abundance||High density|
|Male size range||Gametophyte ca 0.01 mm|
|Male size at maturity|
|Female size range||Gametophyte ca 0.01 mm|
|Female size at maturity|
|Growth form||Arborescent / Arbuscular|
|Characteristic feeding method||Autotroph|
|Typically feeds on||Not relevant|
|Is the species harmful?||No|
The adult plant exhibits no gender but the gametophytes are dioecious. The approximate size of male and female gametophytes are given.
The growth rate during maximal growth is reported. Adults grow rapidly until about 5 years old. Peak growth occurs during winter (November to June) and stops in summer initiated by a photoperiodic response to day length. The total carbon content of canopy lamina is reported to vary with season reflecting a change in carbohydrate storage (Sjøtun, 1996). Carbon content is high in the summer and autumn and starts to decrease in winter with the onset of growth. The old blade is replaced by a new blade formed between the meristem (top of stripe) and the old blade. Nutrients from the old blade contribute to growth. The old blade is shed in spring to early summer.
In Laminaria hyperborea, the proportion of growth allocated to various regions of the plant is reported to vary with both the age of the plant and its habitat (Sjøtun & Fredriksen, 1995). The proportion of growth allocated to the stipe and hapteron, for instance, increases with exposure, the latter probably helping the plant to remain attached and help it to survive in exposed localities (Sjøtun & Fredriksen, 1995). In one year old plants however, growth mainly occurred in the lamina in order to maximize the area for photosynthesis in the light limited understorey.
|Physiographic preferences||Open coast, Strait / sound, Ria / Voe, Enclosed coast / Embayment|
|Biological zone preferences||Lower infralittoral, Upper infralittoral|
|Substratum / habitat preferences||Artificial (man-made), Bedrock, Cobbles, Large to very large boulders, Pebbles|
|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, Moderately exposed, Very exposed|
|Salinity preferences||Full (30-40 psu)|
|Other preferences||No text entered|
|Migration Pattern||Non-migratory / resident|
|Reproductive type||Gonochoristic (dioecious)|
|Reproductive frequency||Annual episodic|
|Fecundity (number of eggs)||>1,000,000|
|Generation time||2-5 years|
|Age at maturity||2 -6 years|
|Season||September - April|
|Life span||11-20 years|
|Larval/juvenile development||Spores (sexual / asexual)|
|Duration of larval stage||See additional information|
|Larval dispersal potential||0 - 10 km|
|Larval settlement period||Can be all year round (see additional information)|
Laminaria hyperborea is a perennial and lives for up to 20 years. Longevity is thought to be higher in its northern distribution (Sjøtun et al., 1993). Spores are produced from sori over most of the blade surface (except most distal or proximal areas) over 6-7 weeks in winter (September - April) (Kain, 1975). Most young sporophytes (germlings) appear in spring but can appear all year round depending on conditions (Birkett et al., 1998b).
Laminarians exhibit alternation of generations and morphologically distinct reproductive phases. The obvious plant is the sporophyte (diploid) producing vast numbers of meiotic haploid zoospores from 'sori'. The flagellated zoospores are about 5 microns in diameter (Sauvageau, 1918; cited in Kain, 1979) and may be transported at least 5 km from the parent (Jónsson, 1972, cited in Norton, 1992). They lose their flagella after 24 hrs (Kain, 1964) and settle on the available substrata. However, settling rate is dependant on the local currents, therefore larval settling time is probably longer than 1 day (Fredriksen et al., 1995). The zoospores develop into microscopic dioecious gametophytes. These become fertile in 10 days in optimal conditions.
Male gametophytes release motile sperm that fertilize eggs of female gametophytes and the resultant zygote develops into the new sporophyte. Mass and rapid sperm release was initiated by adding a drop of sea water, into which female Laminaria hyperborea gametophytes had released eggs, to the male gametophyte culture medium, suggesting the eggs produce pheromones which induce the release of and attract the sperm (Lüning & Müller, 1978).
Maturation of the gametophytes can be delayed under less optimal conditions and development remains vegetative. For example, Lüning (1980) reported that fertility, the induction of fertilization in male and female gametophytes, depended on a critical quantum dose of blue light. Fragments of damaged vegetative gametophytes may develop into separate gametophytes (only a few cells are required) hence reproductive potential may be increased. If optimal conditions return the gametophyte may become fertile and produce gametes. Spore production may be inhibited by epifauna such as Membranipora membranacea (sea mat) and endophytes such as Streblonema sp. (Kain, 1975b).
The MarLIN sensitivity assessment approach used below has been superseded by the MarESA (Marine Evidence-based Sensitivity Assessment) approach (see menu). The MarLIN approach was used for assessments from 1999-2010. The MarESA approach reflects the recent conservation imperatives and terminology and is used for sensitivity assessments from 2014 onwards.
|Removal of the substratum would entail removal of the plants themselves, juvenile sporophytes (germlings) and gametophytes. They can not re-attach once removed and would be swept away. Experimental clearance experiments (Kain, 1979) in the Isle of Man showed that Laminaria hyperborea out-competed opportunistic species (e.g. Alaria esculenta, Saccorhiza polyschides and Desmarestia spp.) and returned to near control levels of biomass within 3 years at 0.8 m but that recovery was slower at 4.4m. However, Kain (1979) noted that grazing would slow recovery since, even though they did not prevent spore settlement, few sporophytes survived after 1 year in the presence of Echinus esculentus. These experiments did not remove the gametophyte 'seed' bank. Research on harvested populations of Laminaria hyperborea in Norway suggests that kelp forest biomass returned to pre-harvesting levels after 1-2 years, but that the plants were mainly small (1m) and that the age structure of the population was shifted towards younger plants. Sivertsen (1991, cited in Birkett et al., 1998b) showed that kelp populations stabilize after about 4-5 year post-harvesting. Re-growth was due primarily to growth of viable juveniles after harvesting. Current advice in Norway suggest that kelp forest should be left for 7-10 years after harvesting for the kelp biomass and non-kelp species to recover (Birkett et al., 1998b). Therefore, recovery is dependant on the depth (light availability) and grazing. However, given the potentially large number of spores and gametophytes it is likely that recolonization would occur rapidly and sporophytes may grow up to 0.94 cm /day under optimal conditions.|
|Although smothering of the adult sporophyte may reduce photosynthetic activity it is unlikely to cause damage. However, juvenile sporophytes may be smothered and their growth inhibited. The germlings, zoospores and gametophytes are likely to be intolerant of smothering.|
|Increased sedimentation may result in smothering of adults (sporophytes), germlings and gametophytes (see above). It may also prevent spore attachment (J. Jones, pers. comm.). Increased sediment deposition may increase sediment scour. However, the most likely effect of increased siltation will be increased light attenuation and turbidity (see below).|
|Laminaria hyperborea is primarily a subtidal species and is unlikely to experience desiccation except during extreme low waters events. It is likely to be highly intolerant of desiccation, and should the single meristem (growth region) be destroyed the plant will die. Although individuals at the top of the shore may be lost the majority of population is found subtidally and is unlikely to be affected.|
|Laminaria hyperborea is primarily a subtidal species and is likely to be highly intolerant of increases in emergence. Its upper limit on the shore is in part dependant on the emergence regime as well as competition from more tolerant species such as Laminaria digitata. An increase in emergence time is likely to depress its upper limit on the shore.|
|The morphology of the stipe and blade vary with water flow rate. 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. It is usually absent from areas of high wave action or strong currents although in Norway it can do well in rapids (J. Jones, pers. comm.).|
|Birkett et al. (1998) suggest that kelp are stenothermal (intolerant of temperature change) and that upper and lower lethal limits for kelp would be between 1-2 °C above or below the normal temperature tolerances. The optimum temperature for the development of Laminaria hyperborea gametophytes and young sporophytes is between 10-17 °C (Kain, 1971). Above 17 °C, gamete survival is reduced (Kain, 1971) and gametogenesis is inhibited at 21 °C in this species (tom Dieck, 1992). Given its distribution in the North Atlantic this species is likely to be tolerant of low temperatures. This species is likely to be intolerant of change in temperature equivalent to either benchmark outside its normal range. 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).|
|The light penetration influences the maximum depth at which kelps species can grow. Dring (1982) reported that laminarians grow at depths at which the light levels are reduced to 1 percent of incident light at the surface. This varies with the turbidity of the sea water from 100 m in the Mediterranean to only 6-7 m in the silt laden German Bight to a maximum of about 35 m in Atlantic European waters. In very turbid waters the depth limit for kelp may be limited to 2 m or it may be absent completely, e.g. Severn Estuary) (Birkett et al., 1998b; Lüning, 1990). Increased turbidity due to coastal engineering, dredging, cooling water plumes have been reported to result in the loss of local kelp forest. Suspended material in vicinity of sewage outfalls have been reported to result in reduced the depth range and the fewer new plants under the canopy. The quality or wavelength of light also affects kelps. Red light favours the accumulation of carbohydrates and blue light enhances protein synthesis, enzyme activity, respiration and is important for the formation of oogonia (eggs) in gametophytes (Dring, 1988). Dissolved organic materials (yellow substance or gelbstoff) absorbs blue light strongly, therefore changes in riverine input or other land based runoff are likely to influence kelp density and distribution. Laminaria hyperborea is likely to be intolerant of a increase of light attenuation of 30 percent of incident surface illumination but would probably not be destroyed within 5 weeks. However, it is likely to be highly intolerant of an increase in turbidity for longer periods especially in deeper waters.|
|Laminaria hyperborea is unable to survive where wave action is extreme because of its large frond area attached to a stiff stipe which is liable to snap. Wave action depresses the upper limit of populations to several metres below low water. In Norway, for example, the upper and lower limits of Laminaria hyperborea are raised from 5 to 0 m and 32 to 26 m from exposed to sheltered sites respectively (Kain, 1971b). It is absent from Rockall possibly due to extreme exposure and strong currents or geographical isolation. Older and larger plants, especially if the holdfasts are weakened by feeding by Helcion pellucidum, are most intolerant of wave action and populations affected by wave action have a reduced age range. As wave exposure increases Laminaria hyperborea is out-competed by Laminaria digitata or Alaria esculenta. In a study in Norway (Sjøtun et al., 1993) Laminaria hyperborea from the most wave exposed site (in Finnmark) exhibited the lowest annual biological productivity per plant. Furthermore, of the four most exposed sites, three of them corresponded with the lowest mean standing crops (fresh weights). Therefore, Laminaria hyperborea is probably highly intolerant of increases in exposure at the benchmark level. It could benefit from decreases in wave exposure, possibly extending its upper limit up the shore, however this would only happen if its upper limit was depressed below the lowest astronomical tide (LAT) as it is highly intolerant of emergence (J. Jones, pers. comm.).|
|Tolerant||Not relevant||Not sensitive||Not relevant|
|Plants have no known sound or vibration receptors|
|Tolerant||Not relevant||Not sensitive||Not relevant|
|Macroalgae are not known to react to the rapid changes in light and shade that would be associated with movement and have no known visual receptors.|
|Physical disturbance caused by a scallop dredge or equivalent impact is likely to have similar effects to that of harvesting, although not so severe (see below). Plants are likely to be removed or damaged by a passing dredge. Therefore, an intolerance of intermediate has been recorded.. Recovery is likely to moderate.|
|Laminaria hyperborea cannot re-attach once removed and would be swept away. Experimental clearance experiments (Kain, 1979) in Isle of Man showed that Laminaria hyperborea out competed other opportunistic species (e.g. Alaria esculenta, Saccorhiza polyschides and Desmarestia spp.) and returned to near control levels of biomass within 3 years at 0.8m but that recovery was slower at 4.4m. However, Kain (1979) noted that grazing would slow recovery since, even though they did not prevent spore settlement, few sporophytes survived after 1 year in the presence of Echinus esculentus. These experiments did not remove the gametophyte 'seed' bank. Research on harvested populations of Laminaria hyperborea in Norway suggests that kelp forest biomass returned to pre-harvesting levels after 1-2 years, but that the plants were mainly small (1m) and that the age structure of the population was shifted towards younger plants. Sivertsen (1991, cited in Birkett et al., 1998) showed that kelp populations stabilize after about 4-5 year post-harvesting. After 4 years post harvesting, kelps had only two thirds of their pre-harvesting canopy height. Re-growth was due primarily to growth of viable juveniles after harvesting. Current advice in Norway suggest that kelp forest should be left for 7-10 years after harvesting for the kelp biomass and non-kelp species to recover (Birkett et al., 1998b). Therefore, recovery is dependant on the depth (light availability) and grazing. However, given the potentially large number of spores and gametophytes it is likely that recolonization would occur rapidly and sporophytes may grow up to 0.94 cm /day under optimal conditions.|
|Hopkin & Kain (1978) examined the effect of Cu, Zn, Hg and Cd on Laminaria hyperborea gametophytes and sporophytes. Sublethal effects on sporophyte development, growth and respiration were shown at concentrations higher than the short term benchmark for Hg, Zn and Cd. Hg was found to be lethal at 0.05 mg/l. However, Cu affected sporophyte development at 0.01mg/l, lower than the benchmark level but was lethal at 0.1 mg/l. However, this report did not examine other heavy metals or their synergistic effects.|
|Mucilaginous slime coating kelp fronds is thought to protect them from coatings of oil. Hydrocarbons in solution reduce photosynthesis and may be algicidal. Reduction in photosynthesis depends on the type of oil, its concentration, length of exposure, method used to prepare oil-water mixture and irradiance in experimental trials (Lobban & Harrison, 1994). The sublittoral fringe populations of Laminaria hyperborea would be most vulnerable to oiling. Subtidal populations being only exposed to dispersed oil or oil adsorbed to particles. Kelps are relatively insensitive to dispersants (Birkett et al., 1998). Three days exposure to 1 percent diesel emulsion reduced photosynthesis completely in young Macrocytsis plants. Laminaria digitata exposed to diesel oil at 130 microgrammes per litre reduced growth by 50 percent in a two year experiment. No growth inhibition was noted at 30 microgrammes per litre and the plants recovered completed in oil-free conditions. Holt et al. (1995) report that oil spills in the USA and the Torry Canyon spillage had little effect on kelp forest. Respiration in Laminaria hyperborea was inhibited by phenol at 100 mg/l (100 ppm).|
|No information||No information||No information||Not relevant|
|All kelp species are efficient absorbers of nutrients (nitrates and phosphates) and can take up and store excess nutrients. Dring (1982) reports that storage of nitrates in winter (when nutrients are plentiful) allows Laminarians to continue growth for 2-3 months after the spring decrease in sea nutrients levels. Although growth is negligible in summer, photosynthesis remains high and reserves of carbohydrates are built up. These carbohydrate reserves peak in autumn, are translocated to the meristem in winter and allow rapid growth in winter when nutrient levels are high. 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. However, eutrophication is associated with loss of perennial macrophytes, a reduction in the depth range and replacement by mussels or opportunistic algae species (Fletcher, 1996; Birkett et al., 1998b) presumably due to indirect effects such as increased turbidity. Increased nutrients may increase growth of epiphytes and plankton, resulting in reduced light penetration for photosynthesis and a subsequent reduction in the depth at which kelp could grow and possibly competition with juvenile sporophytes. Therefore, a rank of intermediate intolerance has been given to represent the likely indirect effects on turbidity and competition.|
|Lüning (1990) suggests that kelps are stenohaline and that their tolerance to salinity covers a range between 16 - 50 psu. Optimal growth probably occurs between 30 -35 psu and grow rates are likely to be affected by periodic salinity stress. Hopkin & Kain (1978) stated that early sporophytes of Laminaria hyperborea grew optimally between 20 -35 psu but did not survive at 6 psu. Birkett et al. (1998) suggest that long term changes in salinity may result in loss of affected kelp beds.|
|No information||Not relevant||No information||Not relevant|
|Little information on the effects of oxygen depletion on macroalgae was found. Kinne (1972) reports that reduced oxygen concentrations inhibit both photosynthesis and respiration. The effects of decreased oxygen concentration equivalent of the benchmark would be greatest during dark when the kelps are dependant on respiration.|
|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.|
|No information||Not relevant||No information||Not relevant|
|The Japanese kelp Undaria pinnatifida (wakame) has recently spread to the south coast of England from Brittany where it was introduced for aquaculture. It is presently restricted to man made structures but could spread in ballast water of commercial or recreational boats and shipping. Its potential competition with other kelps in the UK, including Laminaria hyperborea requires further study (Birkettet al., 1998).|
|Research on harvested populations of Laminaria hyperborea in Norway suggests that kelp forest biomass returned to pre-harvesting levels after 1-2 years, but that the plants were mainly small (1m) and that the age structure of the population was shifted towards younger plants. Sivertsen (1991; cited in Birkett et al., 1998) showed that kelp populations stabilize after about 4-5 year post-harvesting. Re-growth was due primarily to growth of viable juveniles after harvesting. Current advice in Norway suggest that kelp forest should be left for 7-10 years after harvesting for the kelp biomass and non-kelp species to recover (Birkett et al., 1998). Therefore recovery is dependant on the depth (light availability) and grazing. However, given the potentially large number of spores and gametophytes it is likely that recolonization would occur rapidly and sporophytes may grow up to 0.94 cm /day under optimal conditions. Evidence from storm damage indicates that kelp forest can regrow within 14 months. Experimental clearance experiments (Kain, 1979) in Isle of Man showed that Laminaria hyperborea out competed other opportunistic species (e.g. Alaria esculenta, Saccorhiza polyschides and Desmarestia spp.) and returned to near control levels of biomass within 3 years at 0.8 m but that recovery was slower at 4.4 m. However, Kain (1979) noted that grazing would slow recovery since, even though it did not prevent spore settlement, few sporophytes survived after 1 year in the presence of by Echinus esculentus. These experiments did not remove the gametophyte 'seed' bank.|
|Removal of urchin predators such as lobsters or crawfish has been implicated in increases in urchin populations and therefore the creation of 'urchin barrens' and the loss of kelp beds (Birkett et al., 1998b). 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, the evidence is equivocal as sea urchin barrens occur in areas where lobsters are not found (Birkett et al., 1998; Hawkins & Raffaelli, 1999). It is likely that there is a complex interaction between sea urchin recruitment and predation. However, removal of predators or other grazers may perturb the ecosystem making it more intolerant of natural fluctuations in sea urchin numbers or other perturbations.|
- no data -
|National (GB) importance||-||Global red list (IUCN) category||-|
|Origin||-||Date Arrived||Not relevant|
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This review can be cited as:
Last Updated: 03/07/2007