Laminaria digitata, ascidians and bryozoans on tide-swept sublittoral fringe rock
Ecological and functional relationships
Kelp habitats are dynamic ecosystems where competition for space, light and food result in patchy distribution patterns of flora and fauna. Kelp beds are diverse species rich habitats and over 1,200 species have been recorded in UK moderately exposed kelp biotopes (MIR.KR) (Birkett et al., 1998b). Kelps are major primary producers; up to 90% of kelp production enters the detrital food web and is probably a major contributor of organic carbon to surrounding communities (Birkett et al., 1998b). Major interactions are thought to be the effects of competition for space, shading, herbivory and predation.
In most kelp biotopes there is evidence of strong competition for space, especially for space on a favourable substratum. Competition may occur between individual plants of the same species, between kelps and substratum-colonizing species of animals and other algae and between colonial animals and encrusting algae. Competition for space between individuals and species is dynamic, resulting in a constantly changing patchwork of species covering any suitable substrata within the biotope, including the surface of the kelp plants themselves. This is especially true of the components of tide-swept biotopes such as £MIR.Ldig.T£. Tide swept biotopes offer luxuriant conditions for suspension feeders by providing a continual supply of food and removing finer sediment that may otherwise interfere with their delicate feeding apparatus. As a result, strong competition between the suspension feeders that thrive in this biotope will mean that any available substratum is likely to be colonized. Much of the rock surface will be covered by a 'foundation' of encrusting calcareous algae on top of which other species will grow.
The blades of Laminaria digitata plants form a canopy layer, which may cut off much of the incident irradiance. This restricts the development of species with high light demands so that the understorey of plants becomes dominated by shade tolerant red algae including Corallinaceae, Palmaria palmata, Chondrus crispus and Ceramium nodulosum. It also allows species normally restricted to the lower infralittoral in kelp-free areas to compete more effectively in the reduced light levels of the kelp bed and so are found at shallower depths.
Within kelp beds there are relatively few species that graze either the kelp or the understorey algae directly, as the enzymes required to directly utilize algae as food are not common. However, the gastropod Gibbula cineraria is frequently found in this biotope and may graze the kelp, foliose red seaweeds and the rock below. The blue-rayed limpet Helcion pellucidum also grazes on kelp and, when younger, red seaweeds such as Mastocarpus stellatus, which is commonly found in the understory of this biotope. The edible sea urchin Echinus and green sea urchin Psammechinus milaris also graze on kelp species in addition to prey species such as bryozoans, tunicates and hydroids.
Predation within kelp beds has not been well studied in the United Kingdom and very little is known of the predator-prey relationships for the many species occurring in kelp beds. The common shore crab Carcinus maenas is probably the largest mobile predator associated with this biotope and preys upon Gibbula cineraria.
As mentioned previously, tide swept biotopes offer a continual supply of suspended particulate matter that support a thriving suspension feeding community. Suspension feeders in MIR.Ldig.T represent several different phyla.Sponges Of the sponges, the breadcrumb sponge Halichondria panicea is most commonly associated with this biotope. This species is usually found as an encrusting mat on rock and algae. Hymeniacidon perleve is also likely to be present.
Ascidians Both solitary and colonial ascidians are found in this biotope. The colonial ascidians Botryllus schlosseri (the star ascidian) and Botrylloides leachi, and the solitary baked bean sea squirt Dendrodoa grossularia are all frequent.
Cnidaria Several hydroid species are commonly found on rock below the kelp in this biotope, especially Dynamena pumila and Sertularia argentea.
Crustacea Crustacean suspension feeders associated with this biotope are not the most important group, in terms of frequency and abundance, but include the barnacles Balanus crenatus and Semibalanus balanoides.
Annelida The tube worm Pomatoceros triqueter is the most common suspension feeing annelid associated within this biotope. It was found in two thirds of the records of this biotope and can rapidly colonize patches of bare rock. Spirorbid worms may be found.
Bryozoa Alcyonidium gelatinosum, Alcyonidium hirsutum, Electra pilosa, Membranipora membranacea and Scrupocellaria spp. are all likely to compete for space on the fronds and stipes of the kelp plants.
The dominance of suspension feeding fauna indicates the importance of planktonic input to the benthic community of the biotope. Although very little information is available about planktonic communities in kelp beds it can be assumed that there will be larger inputs of larval stages from species with bentho-pelagic life cycles than in the general plankton (Birkett et al., 1998b).
Kelp plants are also exploited as a habitat; the holdfast, stipe and frond each support a different type of community, although only the oldest Laminaria digitata plants will have epiphytic flora and fauna on the stipe (which is smooth in all but the oldest individuals). However, holdfasts shelter a particularly rich diversity of animals from a wide range of taxa (see Habitat Complexity). Epiphytes on the stipe may include the sponge Halichondria panicea and red algae Palmaria palmata and Phycodrys rubens whereas the frond is more likely to be colonized by the bryozoan Membranipora membranacea.
Seasonal and longer term change
Present understanding of the natural fluctuations in the species assemblages, populations, distribution and diversity of species in kelp beds is limited. The plants in this biotope are likely to experience some seasonal change in abundance, the general pattern being a lower percentage cover over the winter months. However, this biotope is limited to extremely sheltered habitats and therefore, the occurrence of winter storms is unlikely to affect it to the same extent that more wave exposed habitats would be affected.
- Growth rate of Laminaria digitata is seasonally controlled with a period of rapid growth from February to July and one of slower growth from August to January. Increased wave exposure and storms experienced during winter months may erode Laminaria digitata blades and reduce the overall standing biomass. Periodic storms may remove older and weaker plants creating patches cleared of kelp. Cleared patches may encourage growth of sporelings or gametophyte maturation. Growth of understorey algae may also be reduced in the winter months.
- Concomitant with the reduction in available surface area of Laminaria digitata blades, a proportion of epiphytic bryozoans, ascidians and sponges will also be lost. However, epilithic representatives of these species will remain on the bedrock and boulders.
- Increased wave exposure and storm frequency over the winter months may also increase the frequency of impacts from wave driven debris, such as pebbles and boulders. These impacts may create 'bare' patches on the surface of the bedrock, and the boulders themselves, which may be colonized by fast growing species including the tube worm Pomatoceros triqueter.
Habitat structure and complexity
Owing to the tide-swept habitat with which this biotope is associated, a diverse marine life is supported. The fast currents provide a continual supply of suspended material sustaining a profusion of both active and passive suspension feeders that dominate the fauna. Fine sediment is removed by the current and the settlement of material, that could otherwise be detrimental to the suspension feeders, is prevented. It is the complex structure of this habitat and its many different niches that allow such a diverse range of suspension feeders to coexist. Almost every possible substratum including the bedrock, boulders and cobbles, and the holdfast, stipe and blade of the Laminaria digitata
itself, is covered with various flora and fauna. In addition to the luxuriant conditions for suspension feeders, Hiscock (1983) lists some the benefits of strong water movement to include the potential for a greater photosynthetic efficiency, thereby possibly increasing the depth penetration of the algae. Increased water movement has been associated with an increase in photosynthesis in several algal species including Fucus serratus
and Ascophyllum nodosum
(Robbins, 1968, cited in Hiscock, 1983).
- Holdfasts provide refuge to a wide variety of animals supporting a diverse fauna that may include polychaetes, small crabs, gastropods, bivalves, and amphipods.
- Kelp fronds are likely to be colonized by encrusting bryozoans (e.g. Membranipora membranacea), ascidians (e.g. Botryllus schlosseri), hydroids (e.g. Dynamena pumila) and sponges (e.g. Halichondria panicea).
- Stipes of Laminaria digitata can support a considerable epiphytic flora, mainly of smaller species (Gayral & Cosson, 1973; Jones et al., 2000).
- The bedrock and boulders offer surfaces for settlement and shelter of species and are colonized by encrusting and foliose red algae but dominated by animals including ascidians, bryozoans, sponges and tubicolous worms.
- Kelp plants are major primary producers in shallow rocky marine habitats in Britain and Ireland. Within the euphotic zone, kelps produce nearly 75% of the net carbon fixed and large kelps often produce annually well in excess of a kilogram of carbon per square metre of shore. However, only about 10% of this productivity is directly grazed. Kelps contribute 2-3 times their standing biomass each year as particulate detritus and dissolved organic matter that provides the energy supply for filter feeders and detritivores in and around the kelp bed. Dissolved organic carbon, algal fragments and microbial film organisms are continually removed by the sea, which may enter the food chain of local subtidal ecosystems, or be exported further offshore. The Corallinaceae and foliose red algae, although not as significant as the kelp, also contribute to primary production within this biotope.
- The fast currents associated with this biotope provide a continual supply of suspended material that sustains a diverse suspension feeding community. Suspension feeders including sponges, bryozoans, ascidians and hydroids, represent the dominant fauna in this biotope highlighting the importance of secondary production.
- Rocky shores make a contribution to the food of many marine species through the production of planktonic larvae and propagules which contribute to pelagic food chains.
- Laminaria digitata plants are fertile all year round with maximum production of spores in July - August and November - December. Young sporophytes (germlings) appear all year with maxima in spring and autumn. Chapman (1981) demonstrated that substantial recruitment of Laminaria digitata plants to areas barren of kelp plants was possible up to 600 m away from reproductive plants.
- Kelp plants themselves can affect recruitment in other species through their influence on the underlying substrata. Shading and mechanical sweeping, for example, will adversely affect settling larvae and post settlement survival.
- With respect to the underlying red algae, tetrasporangia from Corallina officinalis have been recorded throughout the year although settlement occurs after a couple of days which has the potential to limit dispersal. Recruitment in dulse, Palmaria palmata, is most certainly limited in terms of dispersal. Females do not release carpospores so male gametophytes produce spermatia which sink rapidly to enable the male and females gametes to come into contact for fertilization. Lithophyllum incrustans reproduce annually and it has been calculated that 1 mm² of reproductive thallus produces 17.5 million bispores per year with an average settlement of only 55 sporelings/year (Edyvean & Ford, 1984).
- The majority of characteristic fauna associated with this biotope produce planktonic larvae and therefore, depending on respective plankton durations, recruitment is possible from both local sources and populations further away. Breeding in the bryozoan Membranipora membranacea continues through early summer with planktonic cyphonautes settling proceeding into early autumn (Ryland & Hayward, 1977). Pomatoceros triqueter, a tubeworm, produces planktonic all year around, although settlement appears to be limited in winter months.
Time for community to reach maturity
Kain (1975) examined the recolonization of cleared concrete blocks by kelp plants and other algae and found that Laminaria digitata
plants were re-established within 2 years and that red algae returned with a year. Many other characterizing species have planktonic larvae and/or are mobile and so can migrate into the affected area. Colonization of most species of fauna inhabiting kelp holdfast, for example, were found as early as one year after kelp trawling of Laminaria hyperborea
plants in Norway, although numbers of both individuals and species, especially isopods and amphipods, increase with a corresponding increase in holdfast size (Christie et al.
, 1998). However, although these species colonize the biotope quite rapidly maturity of the overall community is likely to be longer (see 'Recoverability'). For example, encrusting coralline algae such as Lithophyllum incrustans
are slow growing (2-7 mm per annum - see Irvine & Chamberlain, 1994) and recruitment of other species to the kelp bed may take longer. In dredged kelp beds in Norway for example, although the rock between Laminaria hyperborea
plants was uniformly covered with coralline algae after 3 years, the more diverse community of cnidarians, bryozoans and sponges associated with coralline algae seen on undredged plots was absent (Rinde et al.
, 1992, cited in Birkett et al.
, 1998). Although it was suggested that the kelp forest recovered to an almost 'normal' state within 3 to 4 years, full biological restoration after harvesting may take at least ten years (Birkett et al
This review can be cited as follows:
Laminaria digitata, ascidians and bryozoans on tide-swept sublittoral fringe rock.
Marine Life Information Network: Biology and Sensitivity Key Information Sub-programme [on-line].
Plymouth: Marine Biological Association of the United Kingdom.
Available from: <http://www.marlin.ac.uk/habitatecology.php?habitatid=273&code=1997>