Fucus serratus and under-boulder fauna on exposed to moderately exposed lower eulittoral boulders

24-02-2005
Researched byDr Keith Hiscock Refereed byDr Bob Foster-Smith
EUNIS CodeA1.2142 EUNIS NameFucus serratus and under-boulder fauna on exposed to moderately exposed lower eulittoral boulders

Summary

UK and Ireland classification

EUNIS 2008A1.2142Fucus serratus and under-boulder fauna on exposed to moderately exposed lower eulittoral boulders
EUNIS 2006A1.2142Fucus serratus and under-boulder fauna on exposed to moderately exposed lower eulittoral boulders
JNCC 2004LR.MLR.BF.Fser.BoFucus serratus and under-boulder fauna on exposed to moderately exposed lower eulittoral boulders
1997 BiotopeLR.MLR.BF.Fser.Fser.BoFucus serratus and under-boulder fauna on lower eulittoral boulders

Description

Underboulder communities are found from midshore downwards on moderately exposed to sheltered boulder shores. The community present under boulders is of the character described here where the boulders are clear of sediment, in pools and/or on open rock from the lower midshore downwards. Characteristic species include the hairy porcelain crab Porcellana platycheles, the long-clawed porcelain crab Pisidia longicornis and juvenile edible crabs Cancer pagurus. Also present beneath the boulders are often high densities of the barnacle Balanus crenatus, the keel worm Pomatoceros spp., spirorbid worms, gammarid amphipods and a few small gastropods and mussels. The encrusting bryozoans Umbonula littoralis and Schizoporella unicornis and encrusting colonies of the sponges Hymeniacidon perleve and Halichondria panicea and the star ascidian Botryllus schlosseri are also typical of this habitat. The richest examples also contain a variety of brittlestars, ascidians and small hydroids. (Biotope description adapted from Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997b).

Recorded distribution in Britain and Ireland

All around Britain but known to be particularly well developed along the coast of Northumberland, in the Plymouth area, in the Isles of Scilly, in the Menai Strait (North Wales) and in tidal rapids at the entrances of sea lochs, Hebridean obs and (in Ireland) loughs.

Depth range

-

Additional information

MLR.Fser.Fser.Bo is also representative of MIR.Ldig.Ldig.Bo where the underboulder fauna is very similar.
Underboulder communities are entirely different from the communities present on the tops and sides of boulders in this biotope.

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Further information sources

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JNCC

Habitat review

Ecology

Ecological and functional relationships

  • The fauna are dominated by a variety of active and passive suspension feeders such as encrusting sponges (e.g. Halichondria panicea), solitary ascidians (e.g. Dendrodoa grossularia), barnacles (e.g. Balanus crenatus), spirorbid worms (e.g. Spirorbis spirorbis), hydroids (e.g. Dynamena pumila), bryozoans (e.g. the sea mat Electra pilosa and the encrusting bryozoan Umbonula littoralis) and colonial ascidians (e.g. Botryllus schlosseri). There is likely to be competition for space between many of the encrusting species. Gordon (1972) noted that competition between bryozoans and ascidians always favoured ascidians. He also noted that Halichondria sponges, even after they had died, prevented the spreading and survival of bryozoans trying to overgrow them.
  • Herbivores include the common periwinkle Littorina littorea, the grey top shell Gibbula cineraria, the green sea urchin Psammechinus milaris and, less frequently, the common limpet Patella vulgata. Herbivorous grazers most likely use the underboulder habitat for shelter but emerge from under the boulders to feed.
  • There are few species that prey on other members of the community but, for instance, dog whelks Nucella lapillus may feed on barnacles and the European cowrie Trivia monacha feeds on the star ascidian Botryllus schlosseri. The common shore crab Carcinas maenas is probably the largest mobile predator associated with MLR.Fser.Fser.Bo. It will move between the boulders and pebbles feeding primarily on small molluscs, especially Littorina spp., annelids and other crustacea. It will also consume algal material. Non-mobile carnivores include the beadlet anemone Actinia equina which feeds passively.
  • Some resident mobile species are detritivores such as the hairy porcelain crab Porcellana platycheles and some brittle stars.
  • Several species that occur under boulders gain shelter from insolation and predators when the tide is out but are not an integral part of the community; for instance, blennies, crabs and shrimps e.g. the common prawn Palaemon serratus.
Species diversity and disturbance
Boulder communities are subject to frequent wave-induced disturbance, especially during the winter months as storm and wave energy increases. Due to the varying size of boulders likely to be found in MLR.Fser.Fser.Bo, some boulders will be moved around and turned-over more frequently than others.

Larger boulders remain undisturbed for longer periods of time and, consequently, the community on them is likely to be dominated by a few late successional species. In MLR.Fser.Fser.Bo, large stable boulders may be dominated by a few prolific species such as Dendrodoa and Halichondria (Foster-Smith, pers. comm.). In contrast, small boulders are tossed around regularly and are unlikely to reach a 'climax' community as disturbance is too frequent. The frequency of disturbance determines the interval of time over which recolonization can occur (Sousa, 1985) and small boulders sample the available pool of spores and larvae more often (Sousa, 1979a) and they are likely to be characterized by hardy species capable of rapidly colonizing bare space e.g. barnacles, spirorbid worms and bryozoans. Sousa (1979a) noted that, in an algal dominated boulder field in California, boulders subjected to intermediate disturbance frequencies were usually less dominated than those which are frequently disturbed, and always less dominated than boulders which were seldom disturbed. Furthermore, intermediate boulders remained undisturbed for long enough that several species had become dominant but not so long that species had been competitively displaced, resulting in dominance. In other words, intermediate size boulders are likely to be more diverse in terms of species diversity. For this reason, the species composition under boulders within the MLR.Fser.Fser.Bo classification can vary considerably which can be problematic when assessing sensitivity (see Species Composition).

Seasonal and longer term change

Some species of bryozoans and hydroids demonstrate seasonal cycles of growth in spring/summer and regression (die back) in late autumn/winter, over wintering as dormant stages or juvenile stages (see Ryland, 1976; Gili & Hughes, 1995; Hayward & Ryland, 1998). Many of the bryozoans and hydroid species are opportunists adapted to rapid growth and reproduction (r-selected), taking advantage of the spring/summer phytoplankton bloom and more favourable (less stormy) conditions (Dyrynda & Ryland, 1982; Gili & Hughes, 1995). Henry (2002) reported a drastic decline in Dynamena pumila over the winter months in the Bay of Fundy. Foster-Smith (1989) recorded that many encrusting ascidians increased in abundance by late summer under boulders on the Northumbrian coast.

On the boulder shores with which MLR.Fser.Fser.Bo is associated, the increased storm and wave energy over the winter months are likely to significantly influence both the flora and faunal components of MLR.Fser.Fser.Bo. Many boulders and cobbles will be thrown around creating bare patches in encrusting species, ripping seaweed off the boulders and overturning boulders to the detriment of species previously on top of the boulders which may suffer from anoxia and crushing etc. These species are likely to perish if left under the boulder which will mean that the proportion of 'bare' rock will increase. Over the winter months therefore they may be an increase in opportunistic species such as Pomatoceros triqueter. However, the winter months also giver the late successional species a chance to colonize the rocks as other e.g. algae such as Ulva die back. Sousa (1979b) found that Ulva sp. inhibited the colonization of mid-successional species such as Fucus serratus and that these mid-successional species, in turn, inhibited the recruitment of late-successional species such as Gigartina canaliculata. Therefore, these winter months are important for the development and diversity of the biotope as a whole.

Habitat structure and complexity

  • The epilithic community usually occurs as a single layer although competition between encrusting species may result in overgrowth and smothering.
  • Habitat complexity increases where soft rocks are bored by bivalve molluscs creating holes for other species to nestle.
  • Variation occurs especially in relation to the degree of influence of underlying sediments. Physical complexity is increased where boulders lie on top of other loosely-packed boulder creating interstices whilst siltation under the boulders means that deeper silty layers may support detritus-feeding polychaetes (Foster-Smith, pers. comm.). Faunal diversity on the boulder surface will be decreased where the boulders are embedded or partly in sediment. In contrast, there may be flowing water under some boulders (for instance, overflows from pools or lagoonal habitats draining at low water) which creates rich communities.

Productivity

Insufficient information

Recruitment processes

The majority of important and characteristic species associated with MLR.Fser.Fser.Bo have planktonic larvae which recruit frequently. Recruitment in the important species is summarized below.
  • The breadcrumb sponge Halichondria panicea is likely to have a short, annual season of sexual reproduction. Most sponges are hermaphroditic but cross-fertilization normally occurs. The process may be oviparous, where there is a mass spawning of gametes through the osculum which enter a neighbouring individual in the inhalant current. Fertilized eggs are discharged into the sea where they develop into a planula larva. However, in the majority, development is viviparous, whereby the larva develops within the sponge and is then released. Larvae have a short planktonic life of a few hours to a few weeks, so that dispersal is probably limited.
  • In Botryllus schlosseri, up to eight eggs are produced per zooid. After fertilization and development to a tadpole stage, the tadpole is released and is free swimming for up to 36 hours (Berril, 1950; Berril, 1975). This short planktonic stage therefore limits recruitment to nearby colonies.
  • Ingle (1997) indicated that the eggs of Pisidia longicornis were present from March to August in southern England and from February to September in the Mediterranean. The planktonic larvae and highly mobile nature of this crab mean that this species does not necessarily rely on recruitment from ,local sources. Underboulder areas may be important refuges for young crabs, especially Cancer pagurus.
  • The dispersal phase of Umbonula littoralis is probably brief and larvae probably do not travel far therefore recruitment is dependant on local sources. Embryos were recorded as present in the Plymouth area in June and August (Marine Biological Association, 1957), from October and November on the north-east coast of England (Hastings, 1944) and from September to February in Manx waters (Eggleston, 1969).
  • Balanus crenatus releases planktotrophic nauplii larvae between February and September, with peaks in April and late summer when phytoplankton levels are highest. They pass through six nauplii stages before eventually developing into a cyprid larva which is are specialized for settlement. Peak settlement occurs in April and declines until October. The larvae may not settle for a month after release and therefore, dispersal potential is relatively high as is recruitment from distant sources. Metamorphosis usually takes place within 24 hours of settlement.
  • Although asexual reproduction occurs in many ascidians, reproduction in the baked bean ascidian Dendrodoa grossularia is entirely sexual (Millar, 1954). Millar studied reproduction in Dendrodoa grossularia in two locations (the River Crouch in Essex and the Isle of Cumbrae in the Firth of Clyde) and found that reproduction was bi-polar in nature with one peak in spring and another in late autumn, the spring episode being more intense. The average number of eggs produced per individual (over 7 mm in length) was only ca 25-100 eggs in the Clyde and River Crouch respectively. Furthermore, the eggs are brooded internally until the larval stage is reached thereby compressing the free swimming stage.
  • Time for community to reach maturity

    Settlement panels, which attract similar communities to underboulder habitats, may be fully colonized within about 18 months of being placed into the environment (extrapolated from Sutherland & Karlson, 1977; Todd, 1994). Development of 'mature' communities under boulders is likely to occur within two years and there will be dynamic stability, i.e. composition of the community will remain much the same although individual organisms and colonies will die and be replaced by the same species.

    Additional information

    No text entered

Preferences & Distribution

Recorded distribution in Britain and IrelandAll around Britain but known to be particularly well developed along the coast of Northumberland, in the Plymouth area, in the Isles of Scilly, in the Menai Strait (North Wales) and in tidal rapids at the entrances of sea lochs, Hebridean obs and (in Ireland) loughs.

Habitat preferences

Depth Range
Water clarity preferences
Limiting Nutrients No preference
Salinity
Physiographic
Biological Zone
Substratum
Tidal
Wave
Other preferences Flowing water

Additional Information

The richest underboulder communities develop in wave sheltered locations on stable boulders where the downward facing surfaces are clear of sediment and there is flowing water present e.g. Strangford Lough, Menai Strait and Linne Mhuirich.

Species composition

Species found especially in this biotope

Rare or scarce species associated with this biotope

Additional information

Underboulder communities are especially described in Foster-Smith (1989, 1991), Foster-Smith & Foster-Smith (1987), and Hiscock (1984) and in various reports produced during the MNCR survey of Scottish sea lochs.

Sensitivity reviewHow is sensitivity assessed?

Explanation

The structure of MLR.Fser.Fser.Bo communities is likely to vary greatly between boulders. As a result many of the species chosen as indicative to sensitivity have been listed as 'important other', since other designations suggest the species plays a permanent central role in the biotope. A representative of the groups of organisms most likely to be found in the biotope at a given time has been selected, bearing in mind the difference between frequently disturbed and seldom disturbed boulder communities (see Ecological and Functional Relationships).

The breadcrumb sponge Halichondria panicea and star ascidian Botryllus schlosseri have both been listed as important structural species since they are competitively superior to many encrusting bryozoan species likely to occur in underboulder communities. A solitary ascidian, Dendrodoa grossularia, has also been included since its represents a species that is likely to be found in larger boulders that have undergone considerable succession in terms of community development. Umbonula littoralis has been included as a representative of the various encrusting bryozoans likely to be found in MLR.Fser.Fser.Bo. The acorn barnacle Balanus crenatus, along with bryozoans and colonial ascidians, will be an important early colonizer of the rock surface and is likely to be one of the pioneer species on smaller frequently disturbed boulders. Apart from filter feeders, the dominant trophic group, the long-clawed porcelain crab, Pisidia longicornis scavenger has been listed as important other since it is likely to be found in and around the sediment that under the boulder. The broad-clawed porcelain crab Porcellana platycheles may be more commonly associated with MLR.Fser.Fser.Bo than Pisidia longicornis although more information was available for the long-clawed porcelain crab.

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 otherBalanus crenatusAn acorn barnacle
Important structuralBotryllus schlosseriStar ascidian
Important otherDendrodoa grossulariaBaked bean ascidian
Important structuralHalichondria paniceaBreadcrumb sponge
Important otherPisidia longicornisLong-clawed porcelain crab
Important otherUmbonula littoralisAn encrusting bryozoan

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High High Moderate Major decline Low
Substratum removal will result in the loss of the entire MLR.Fser.Fser.Bo community and, therefore, intolerance has been assessed as high. Although mobile species including the long- and broad-clawed porcelain crabs may survive, they are not, in isolation, representative of MLR.Fser.Fser.Bo. Recoverability is likely to be high (see additional information).
High High Moderate Decline Low
Many of the underboulder species are low-lying encrusting forms that cannot escape smothering and are, therefore, especially vulnerable. Over the course of one month, feeding in suspension feeders is likely to be inhibited as a result of the clogging of the feeding apparatus. In addition, deoxygenation will occur due to the decomposition of smothered matter under the boulder. Balanus crenatus can withstand covering by silt provided that the cirri can extend above the silt layer but smothering by 5 cm of sediment would prevent feeding and could cause death. It is likely that many of the important species including the bryozoans and colonial ascidians will experience mortality and accordingly, intolerance has been assessed as high. However, smothering by sand is part of the natural dynamics of some boulders (Foster-Smith, pers. comm.) and the fact that the majority of underboulder communities are downward facing means that the effects of smothering are likely to be relatively short lived. Recoverability is expected to be high (see additional information). (This assessment is for smothering by sediment - some typical underboulder species can survive overgrowth by other species (c.f. Turner, 1988)).
Intermediate High Low Minor decline Low
Underboulder communities face downwards so that silt is unlikely to settle but may clog the feeding structures of some species such as hydroids, bryozoans and ascidians thereby reducing total ingestion over the benchmark period. Umbonula littoralis for example, is expected to have a limited ability to clear itself of silt. Rich underboulder communities are known to occur in turbid waters, for instance, the Menai Strait. However, increased suspended sediment, in combination with areas of low wave energy or water movement may lead to siltation (see water flow rate) and therefore, intolerance has been assessed as intermediate. Recoverability is likely to be high (see additional information).
Tolerant* High Not sensitive Decline Low
A decrease in suspended sediment is likely to be beneficial to most of the underboulder community. The suspension feeders may become more efficient as there would be fewer inorganic particles to clog and interfere with feeding apparatus. Assuming that the decrease in suspended sediment refers to inorganic particles, a reduction in total ingestion in the suspension feeding community is not expected. Therefore, tolerant* has been assessed.
High High Moderate Major decline Low
Underboulder communities are generally damp due to the fact that they are mostly unaffected by the drying influences of wind and insolation. Furthermore, many underboulder communities are in contact with flowing water. Underboulder species on boulders which are turned so that the undersurface community ceases to be shaded and damp are likely to be killed. The number and diversity of species likely to be killed be dependent on the size of the boulder since larger boulders are less likely to be turned over and so will have more developed community (see Ecology). Small rocks frequently turned over (either through natural energy, e.g. by wave energy, or humans) will have fewer species and species that, nevertheless, are opportunistic species characteristic of disturbed environments. Balanus crenatus, for example, were reported to have a mean survival time of 14.4 hours in dry air (Barnes et al., 1963). The community will eventually re-develop on the new underside and therefore, recoverability is expected to be high (see additional information).
Intermediate High Low Decline Low
A one hour change in the time not covered by the sea for a period of one year is unlikely to adversely affect the majority of the MLR.Fser.Fser.Bo community since the habitat is likely to remain shaded and damp. Mobile species such as Pisidia longicornis and Carcinas maenas, because of their mobility, may be able to escape the effects of increased emergence by crawling to damper areas further down the shore.

On balance, however, MLR.Fser.Fser.Bo has been assessed as being of intermediate intolerance to changes in emergence to reflect the likelihood that species at the limits of their tolerance to emergence might be killed. Recoverability is likely to be high (see additional information).

Tolerant Not relevant No change Very low
A decrease in emergence would reduce the influence of desiccation on the community which would be beneficial to the biotope. However, this benefit may be counteracted by the fact that the more submerged boulders may be subject to increased disturbance through wave energy. Larger boulders previously undisturbed may move around more, potentially leading to an increased species diversity (see Ecology).

On balance, MLR.Fser.Fser.Bo has been assessed as tolerant to a decrease in emergence.

Tolerant* Not relevant Not sensitive* No change Low
The richest underboulder communities develop in areas subject to strong tidal flows and, therefore, at the benchmark level, MLR.Fser.Fser.Bo is likely to be tolerant*.
Intermediate High Low Decline Low
A decrease in strength of tidal flow will lead to loss or reduction in abundance of some species and this would most likely be a result of increased siltation. Species including Pisidia longicornis and Umbonula littoralis thrive in habitats that are in areas of moderate to strong water movement. A decrease in water flow rates where wave action is also weak would be likely to result in mortality in, for example, some bryozoans, colonial ascidians and sponges. This is most likely as a secondary effect from siltation but possibly also due to a reduction in food source. Barnes & Bagenal (1951) found that the growth rate of Balanus crenatus epizoic on Nephrops norvegicus was considerably slower than animals on raft exposed panels and this was attributed to reduced currents and increased silt loading of water in the immediate vicinity of Nephrops norvegicus. Intolerance is, therefore, assessed as intermediate. However, recoverability will be high (see additional information).
Low High Low Minor decline Low
The shaded and damp conditions found in underboulder communities may serve to protect the MLR.Fser.Fser.Bo community from extremes of temperature. Nevertheless, the important species found in this biotope have varying levels of tolerance to changes in temperature at the benchmark level and some species living under boulders are normally subtidal species and may be unable to withstand large changes in temperature.
  • Pisidia longicornis occurs in a wide range of temperature regimes from Norway to Angola and it is unlikely that they would be adversely affected by an increase in temperature at the level of the benchmark.
  • The British Isles are at the centre of geographical range for Umbonula littoralis, Botryllus schlosseri and Halichondria panicea suggesting that colonies are likely to be tolerant of both an increase and decrease in temperature at the benchmark level.
  • Balanus crenatus is a boreal species that is likely to be intolerant of increases in water temperature. In Queens Dock, Swansea, where the water temperature was on average 10 °C higher than average due to the effects of a condenser effluent, Balanus crenatus was replaced by the subtropical barnacle Balanus amphitrite. After the water temperature cooled Balanus crenatus returned (Naylor, 1965). Balanus crenatus was unaffected during the severe winter of 1962-63, when average temperatures were 5 to 6 °C below normal (Crisp, 1964a).
  • Gamete release in Dendrodoa grossularia decreases at 15 degrees and is suppressed at 20 degrees and below about 8-11 degrees (Millar, 1954). It is likely to be sensitive to an increase and decrease in temperature at the benchmark level.
On balance, it is likely that overall intolerance to an increase in temperature will be low.
Low High Low Minor decline Low
The shaded and damp conditions found in underboulder communities may serve to protect the MLR.Fser.Fser.Bo community from extremes of temperature. Nevertheless, the important species found in this biotope have varying levels of tolerance to changes in temperature at the benchmark level and some species living under boulders are normally subtidal species and may be unable to withstand large changes in temperature.
  • Pisidia longicornis were adversely affected by the 1962-63 winter in Britain. Crisp (1964a) records that many hundreds were found dead on the strandline at Oxwich, south Wales. In other locations, they were not found on the shore (although could have migrated offshore).
  • The British Isles are at the centre of geographical range for Umbonula littoralis, Botryllus schlosseri and Halichondria panicea suggesting that colonies are likely to be tolerant of both an increase and decrease in temperature at the benchmark level.
  • Gamete release in Dendrodoa grossularia decreases at 15 degrees and is suppressed at 20 degrees and below about 8-11 degrees (Millar, 1954). It is likely to be sensitive to an increase and decrease in temperature at the benchmark level.
On balance, it is likely that overall intolerance to a decrease in temperature will be low
Tolerant Not relevant Not sensitive No change Moderate
Rich underboulder communities are known to occur in turbid waters, for instance, the Menai Strait. Therefore, it has been suggested that MLR.Fser.Fser.Bo is tolerant to an increase in turbidity at the benchmark level.
Tolerant Not relevant No change Moderate
A decrease in turbidity may stimulate phytoplankton production which would be beneficial to the suspension feeding community associated with MLR.Fser.Fser.Bo. Therefore, it has been suggested that MLR.Fser.Fser.Bo is tolerant to an increase in turbidity at the benchmark level.
Intermediate High Low Decline Low
Many of the species likely to be found in MLR.Fser.Fser.Bo communities are probably tolerant of very wave exposed conditions. However, increases in wave exposure may cause more boulders to become mobile and abrade underboulder communities. Increased mobilization of boulders may result in patches of sponges, bryozoans and barnacles being crushed on impact with other boulders. For example, Umbonula littoralis has a hard calcareous skeleton which is likely to be broken through contact with hard surfaces such as cobbles moving around during storms. Crabs and other fragile mobile species are also at risk from being crushed. Furthermore, many of the stable boulders are fused together by algal growth (especially corallines) and breaking up this matrix would adversely affect the community (Foster-Smith, pers. comm.). The release of sediment between boulders may serve to interrupt suspension feeding (see Suspended Sediment above).

MLR.Fser.Fser.Bo is found on shores ranging from wave sheltered to moderately wave exposed and as a result the communities in the biotope between each of these locations will vary anyway and. Therefore, different sites are likely to have varying tolerances with respect to changes in wave exposure. On balance, MLR.Fser.Fser.Bo has been assessed as being of intermediate intolerance to a change I wave exposure since some species may experience mortality although even frequently disturbed boulders with a few pioneer species may still represent MLR.Fser.Fser.Bo. Recovery is expected to be rapid (see additional information).

Intermediate High Low Decline Low
A decrease in wave exposure may facilitate sedimentation which will smother underboulder species resulting in mortality (see Smothering above).

MLR.Fser.Fser.Bo is found on shores ranging from wave sheltered to moderately wave exposed and as a result the communities in the biotope between each of these locations will vary anyway and. Therefore, different sites are likely to have varying tolerances with respect to changes in wave exposure. On balance, MLR.Fser.Fser.Bo has been assessed as being of intermediate intolerance to a change in wave exposure since some species may experience mortality although even frequently disturbed boulders with a few pioneer species may still represent MLR.Fser.Fser.Bo. Recovery is expected to be rapid (see additional information).

Tolerant Not relevant Not relevant Not relevant Moderate
The characteristic and permanent members of the fauna are invertebrates unlikely to detect or be affected by noise.
Tolerant Not relevant Not sensitive Not relevant Moderate
The characteristic and permanent members of the fauna are invertebrates unlikely to detect or be affected by visual presence.
High High Moderate Decline Moderate
In addition to disturbance caused by wave energy, intertidal boulder communities are often disturbed by, for example, bait collectors, inquisitive school groups and field researchers. Boulders left overturned place the organisms on the now upward facing part of the boulder at great risk of desiccation (see Desiccation above). Furthermore, many stable boulders are fused together by algal growth (especially corallines) and breaking this matrix would be very harmful (Foster-Smith, pers. comm.). Furthermore, this disturbance and habitat degradation could change a stable boulder field to an unstable field on a long-term basis (Foster-Smith, pers. comm.). Movement of the boulder surface against other hard surfaces (for instance, during extreme storm events) is likely to cause significant damage to encrusting fauna that is characteristic of the community. Recoverability is expected to be high (see additional information).
High High Moderate Major decline Moderate
Due to the fact that the majority of species likely to be found in the MLR.Fser.Fser.Bo community are permanently attached to the substratum, displacement will have the same effect as substratum removal and, therefore, intolerance has been assessed as high.

Chemical Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
Intermediate High Low Decline Low
Some members of the community, particularly crustaceans and molluscs, may be intolerant of chemicals that may have an adverse effect on reproduction. Chemicals developed as anti-fouling paints have been developed to counter fouling communities which are similar to underboulder communities. Barnacles have a low resilience to chemicals such as dispersants, dependant on the concentration and type of chemical involved (Holt et al., 1995). Hoare & Hiscock (1974) found that Balanus crenatus survived near an acidified halogenated effluent discharge where many other species were killed, suggesting a high tolerance to chemical contamination. Little information is available on the impact of endocrine disrupters on adult barnacles or on the effects of synthetic chemicals on the other important species. However, intolerance has been suggested as intermediate to reflect the likely effects of antifouling chemicals. Component species generally have planktonic larvae and reproduce frequently so that re-colonization will be rapid, providing the environment is clean of any chemicals that were having an adverse effect ob the community.
Heavy metal contamination
No information Not relevant No information Insufficient
information
Not relevant
Barnacles accumulate heavy metals and store them as insoluble granules (Rainbow, 1987). Pyefinch & Mott (1948) recorded a median lethal concentration of 0.19 mg/l copper and 1.35 mg/l mercury, for Balanus crenatus over 24 hours. Barnacles may tolerate fairly high level of heavy metals in nature, for example they are found in Dulas Bay, Anglesey, where copper reaches concentrations of 24.5 µg/l, due to acid mine waste (Foster et al., 1978). However, insufficient information was available on the remaining important species in MLR.Fser.Fser.Bo and sensitivity has not been assessed.
Hydrocarbon contamination
Intermediate High Low Decline Moderate
Little evidence was found. Ryland & de Putron (1998) found no detectable damage to underboulder faunas during oil pollution in Watwick Bay, Pembrokeshire. However, fresh oil is likely to narcotize and kill Decapoda and some Gastropoda. Component species generally have planktonic larvae and reproduce frequently so that re-colonization will be rapid, providing the environment is clean of any chemicals that were having an adverse effect on the community.
Radionuclide contamination
No information Not relevant No information Insufficient
information
Not relevant
Insufficient
information.
Changes in nutrient levels
Tolerant Not relevant Not relevant No change Low
Underboulder communities occur where nutrient concentrations are high in enclosed coastal areas, for instance, the Menai Strait. The underboulder area is shaded and therefore any risks of thereby reducing the likelihood of smothering by ephemeral green algal species that are likely to flourish in the event of nutrient influx. Tolerant has been suggested.
Tolerant Not relevant Not sensitive No change Low
Underboulder communities occur in full to variable salinity habitats although it might be that higher salinity occurs at the outflow of some basins. At the levels expected, MLR.Fser.Fser.Bo is likely to be tolerant to an increase in salinity.
Intermediate High Low Decline Low
Rich underboulder communities occur in outflows from areas of variable salinity (for instance, in the Menai Strait, North Wales and in sea lochs). The communities therefore have some tolerance to at least short-term reduced salinity. However, some of the component species may be intolerant of reduced salinity. Pisidia longicornis and Umbonula littoralis both occur in full salinity and are likely to be intolerant of an acute reduction in salinity. Other species would be very tolerant. Balanus crenatus, for example, can tolerate salinities down to 14 psu if given time to acclimate (Foster, 1970). On balance, an intolerance of intermediate has been suggested to reflect the possibility that some species may experience some mortality. Component species generally have planktonic larvae and reproduce frequently so that re-colonization will be rapid.
Intermediate High Low Minor decline Very low
Underboulder habitats may be subject to lowered oxygen levels due to restricted water flow in calm periods. Also, organic debris that becomes trapped under the boulders may rot and cause de-oxygenated conditions. Some tolerance of low oxygen levels is therefore expected in some situations. However, the richest underboulder communities occur where water flow is strong and almost continuous and might suffer in de-oxygenated conditions. Component species generally have planktonic larvae and reproduce frequently so that re-colonization will be rapid.

Biological Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
No information Not relevant No information Not relevant Not relevant
Insufficient
information.
No information Not relevant No information Not relevant Not relevant
Insufficient
information.
Low High Low Minor decline Low
Species that are extracted from underboulder communities include edible crustaceans which, as scavengers, are not of key importance in the functioning of the community. None of the important species are likely to be targeted for extraction although the collection of other creatures including crabs and shrimps may result in increased physical disturbance, to the detriment of the community (see Physical Disturbance).
Low High Low Minor decline Low

Additional information

Recoverability
The community associated with MLR.Fser.Fser.Bo will very greatly depending on various factors including the size of boulder, wave exposure and the presence or absence of flowing water under the boulder. In addition, it is difficult to identify a ‘climax’ community per se because the extent of community succession will vary greatly between boulders of different sizes etc. Furthermore, because there are no key functional, structural or characterizing species, any combination of the important species could, theoretically, determine the biotope community. Nevertheless, the recolonization of fauna typically associated with MLR.Fser.Fser.Bo will occur within a year or two and recoverability is expected to be high. However, the development of a mature community characteristic of seldom disturbed boulders dominated by e.g. Halichondria panicea and Dendrodoa grossularia may take longer although many boulders will never mature to this stage.

In the study of recolonization of vertical rock wall in Maine (Sebens, 1986), epifaunal and algal crust species were shown to re-colonize cleared areas quickly. For example encrusting bryozoans, tubeworms, tubicolous amphipods and worms, erect hydroids and bryozoans were reported to cover cleared areas within 1-4 months in spring, summer and autumn (Sebens, 1986). Sebens (1985) reported that Halichondria panicea had reached previous cover within two or more years. It was slow to recolonize the cleared areas, only appearing after about a year, although it is relatively fast growing. Balanus crenatus is another important early colonizer of sublittoral rock surfaces (Kitching, 1937) and it heavily colonized a site that was dredged for gravel within 7 months (Kenny & Rees, 1994).

Importance review

Policy/Legislation

Habitats of Principal ImportanceIntertidal boulder communities
Habitats of Conservation ImportanceIntertidal underboulder communities
Habitats Directive Annex 1Reefs
UK Biodiversity Action Plan PriorityIntertidal underboulder communities

Exploitation

Lobsters and winkles are collected from underboulder habitats for human consumption and crabs and shrimps are also collected, primarily for bait (Foster-Smith, pers. comm.). Both activities are very common in some areas e.g. Northumberland (Foster-Smith, pers. comm.). There is incidental damage to communities by crushing when boulders are returned and much greater damage if boulders are left upturned (see Sensitivity).

Additional information

Field study and general exploration requires advice to return boulders carefully to their original locations after removing mobile species that might be crushed. Bait collectors may require to be restricted in their activities if they fail to take care at important sites.

Boulder habitats are a part of "Reefs" in Annex 1 of the Habitats Directive and also occur in "Large shallow inlets and bays" and in "Estuaries".

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Citation

This review can be cited as:

Hiscock, K. 2005. Fucus serratus and under-boulder fauna on exposed to moderately exposed lower eulittoral boulders. 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. Available from: http://www.marlin.ac.uk/habitat/detail/371

Last Updated: 24/02/2005