Dense Lanice conchilega and other polychaetes in tide-swept infralittoral sand and mixed gravelly sand

Researched byGeorgina Budd Refereed byThis information is not refereed.
EUNIS CodeA5.137 EUNIS NameDense Lanice conchilega and other polychaetes in tide-swept infralittoral sand and mixed gravelly sand


UK and Ireland classification

EUNIS 2008A5.137Dense Lanice conchilega and other polychaetes in tide-swept infralittoral sand and mixed gravelly sand
EUNIS 2006A5.137Dense Lanice conchilega and other polychaetes in tide-swept infralittoral sand and mixed gravelly sand
JNCC 2004SS.SCS.ICS.SLanDense Lanice conchilega and other polychaetes in tide-swept infralittoral sand and mixed gravelly sand
1997 BiotopeSS.IGS.FaS.LconDense Lanice conchilega and other polychaetes in tide-swept infralittoral sand


Where strong tidal streams or wave action and coarse sand occur in the shallow sublittoral, dense beds of Lanice conchilega may occur. Several other species of polychaete also occur as infauna e.g. Scoloplos armiger, Chaetozone setosa and Arenicola marina. The dense Lanice biotope (LGS.Lan) on certain lower shores may be a littoral extension of this biotope. This biotope also appears to have a limited occurrence in some Scottish lagoonal entrance channels and some sea lochs. Overall, there may be more than one entity in this biotope. (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Recorded distribution in Britain and Ireland

Present on all coasts of Britain with the exception of the north east coast of Scotland and south east coast of England. Reported in Ireland at Rosbin Cove in Roaringwater Bay, and along the coast from Kilmichael Point southwards around Cahore Point to Raven Point, County Wexford.

Depth range


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Habitat review


Ecological and functional relationships

  • The hydrodynamic regime and sediment composition and interaction between the two are probably the most significant factors structuring the community rather than biological interactions (Tyler, 1977; Warwick & Uncles, 1980; Elliott et al., 1998).
  • Species in the biotope are predominantly suspension and deposit feeders, and probably little direct interaction occurs between them other than competition for space.
  • Some species feed on both suspended particulates and surface deposits (e.g. the bivalves Fabulina fabula and Abra alba and the tube building polychaete Lanice conchilega. Following laboratory experiments, Buhr (1976) concluded that Lanice conchilega was capable of completely replacing deposit feeding by suspension feeding. The absolute amounts of food retained from suspension feeding and the assimilation efficiencies calculated were in the range typical for obligatory suspension-feeding organisms.
  • Studies of Lanice conchilega aggregations in the Wadden Sea ( Zühlke et al.,1998; Dittmann, 1999; Zühlke, 2001) have shown that tubes built by Lanice conchilega can have significant effects on the distribution, density and diversity of other macrobenthic species and meiobenthic nematodes compared to sites with a lower density of Lanice conchilega or ambient sediment without biogenic structures. The polychaete Harmothoe lunulata occurs in aggregations of Lanice conchilega and is often found inside the polychaetes' tubes, possibly being a commensal associated to Lanice conchilega (Zühlke et al., 1998). Juvenile bivalves (Mya arenaria, Mytilus edulis, Macoma balthica) were more frequent in patches with Lanice conchilega and settled especially on the tentacle crown of the worm tubes. In particular, abundances of predatory polychaetes (Eteone longa, Nephtys hombergii, Hediste diversicolor) were higher (Dittmann, 1999). The increased species diversity and abundance recorded in patches of Lanice conchilega are also known to occur around the tubes of other species of polychaetes (Woodin, 1978).
  • In sand, the primitive sea slug Faction tornatilis preys upon tube building polychaetes. A series of choice experiments suggested that the preferred prey items were the polychaetes Owenia fusiformis and Lanice conchilega (Yonow, 1989).
  • Lanice conchilega constitutes an important prey item for curlew, Numenius arquata, bar-tailed godwit, Limosa lapponica and grey plover, Pluvialis squatarola.
  • The amphipods e.g. Ampelisca and Atylus species are probably epistatic grazers, grazing benthic microalgae from sand grains.
  • Amphipods and the infaunal annelid species in the biotope probably interfere strongly with each other. Adult worms probably reduce amphipod numbers by disturbing their burrows and tubes, while high densities of amphipods can prevent establishment of worms by consuming larvae and juveniles (Olafsson & Persson, 1986).
  • Spatial competition probably occurs between the infaunal suspension feeders and deposit feeders. Reworking of sediment by deposit feeders makes the substratum less stable, increases the suspended sediment and makes the environment less suitable for suspension feeders (Rhoads & Young, 1970). Tube building by amphipods and polychaetes stabilizes the sediment and arrests the shift towards a community consisting entirely of deposit feeders. In the coarse sediments in this biotope the suspension feeding species dominate.
  • Amphipods are preyed upon chiefly by nemertean worms (see McDermott, 1984) and demersal fish (Costa & Elliott, 1991).
  • The abundant infauna are preyed upon by carnivorous polychaetes, e.g. phyllodocids species of Anaitides and Eumida, scale worms (e.g. Harmothoe sp.) and Nephtys hombergii.
  • Asterias rubens, predates the bivalves (Aberkali & Trueman, 1985; Elliott et al., 1998).
  • Crabs, particularly Liocarcinus depurator and Carcinus maenas, are scavengers and predators of molluscs and annelids (Thrush, 1986; Elliott et al., 1998).
  • The hermit crab Pagurus bernhardus, brittlestars (e.g. Ophiura albida) and nemerteans are probably scavengers on detritus and carrion.
  • Gobies (e.g. Pomatoschistus species) and flatfish frequent the biotope to feed upon polychaetes, small crustaceans such as amphipods, cumaceans, small crabs, such predators also nip the siphons of bivalves and tails of polychaetes.

Seasonal and longer term change

  • Temporal changes are likely to occur in the community due to seasonal recruitment processes. For instance, in the German Bight, peak abundance of Fabulina fabula (ca 2000 individuals/m²) occurred in September following the main period of spatfall and then decreased to a minimum in February (ca 500 individuals/m²), at which point settlement began to occur again (Salzwedel, 1979). Similarly, temporal evolution of Fabulina fabula in NW Spain showed well marked annual peaks in autumn (Lopez-Jamar et al., 1995). In the German Bight, spatfall for Magelona mirabilis was heaviest in August/September, and for Echinocardium cordatum spatfall was heaviest in August (Bosselmann, 1989).
  • Temporal variations in species richness and abundance are likely to occur due to seasonal patterns of disturbance, such as storms, harsh winters and oxygen deficiencies (Bosselmann, 1989; Lopez-Jamar et al., 1995). The biotope may also be liable to severe substratum disturbance, such as one in 25 year or one in 50 year storms, which can turn over sediment and completely disrupt the community (Elliott et al., 1998).
  • There may be a spring-neap and winter-summer cycle of erosion and deposition of sediment, altering the biotope extent and reflecting changes in hydrodynamic energy (Dyer, 1998).
  • The water temperature in subtidal sandy habitats may vary over 5-10 °C through the year in British coastal waters depending on depth. The variation may have short term but significant effects on species diversity (Buchanan & Moore, 1986).
  • There is a seasonal variation in planktonic production in surface waters. Increased production by phytoplankton in spring and summer enhanced by increasing temperature and irradiance is followed by phytoplankton sedimenting events which correlate with seasonal variations in the organic content of benthic sediments (Thouzeau et al., 1996). Such variations directly influence the food supply of the deposit feeders and suspension feeders in the biotope.

Habitat structure and complexity

  • Tidal streams in particular are a predominant factor influencing the structure of infralittoral sands, causing constant change in the shape, size and position and owing to the mobile nature of the sandy substratum and scour macrophyte communities do not become established.
  • The habitat can be divided into several niches. The illuminated sediment surface supports a flora of microalgae such as diatoms and euglenoids, together with aerobic microbes and ephemeral green algae in the summer months. The aerobic upper layer of sediment supports shallow burrowing species such as amphipods and small crustacea, whilst the reducing layer and deeper anoxic layer support chemoautotrophic bacteria, burrowing polychaetes (e.g. Chaetopterus variopedatus and Arenicola marina) that can irrigate their burrows, and burrowing bivalves (e.g. Abra alba).
  • In fairly homogeneous soft sediments, biotic features play an important role in enhancing species diversity and distribution patterns (Bandeira, 1995; Everett, 1991; Sebens, 1991). Polychaete dwelling tubes, such as those constructed by Lanice conchilega, provide one of the main habitat structures in the intertidal and subtidal zones. The tubes modify benthic boundary layer hydrodynamincs (Eckman et al., 1981), can provide an attachment surface for filamentous algae (Schories & Reise, 1993) and serve as a refuge from predation (Woodin, 1978) (Zühlke et al., 1998). Other biota probably help to stabilize the substratum. For example, the microphytobenthos in the interstices of the sand grains produce mucilaginous secretions which stabilize fine substrata (Tait & Dipper, 1998). The presence of infaunal polychaetes affects the depth of the oxic sediment layer. Tubes of Lanice conchilega and Arenicola marina can penetrate several tens of centimetres into the sediment. Such burrows and tubes allow oxygenated water to penetrate into the sediment indicated by 'halos' of oxidized sediment along burrow and tube walls. The burrow of Arenicola marina is irrigated (and therefore aerated) by intermittent cycles of peristaltic contractions of the body from the tail to the head end. Lanice conchilega is not known to purposely irrigate its tube since food acquisition occurs through surface deposit and suspension feeding, and respiration takes place via gills positioned outside the tube when the animal is feeding. However, the sand mason periodically withdraws from the surface into its tube for a few seconds, therefore acting as a 'piston pump', exchanging interstitial sediment water with overlying water (Forster & Graf, 1995).
  • Productivity

    Production in the biotope is mostly secondary, dependant upon detritus and organic material. Some primary production comes from benthic microalgae and water column phytoplankton. The microphytobenthos consists of unicellular eukaryotic algae and cyanobacteria that grow in the upper several millimetres of illuminated sediments, typically appearing only as a subtle brown or green shading (Elliott et al., 1998). The benthos is supported predominantly by pelagic production and by detrital materials emanating from the coastal fringe (Barnes & Hughes, 1992). According to Barnes & Hughes (1992) the amount of planktonic food reaching the benthos is related to:
    • depth of water through which the material must travel;
    • magnitude of pelagic production;
    • proximity of additional sources of detritus;
    • extent of water movement near the sea bed, bringing about the renewal of suspended supplies;
    In the relatively shallow waters around the British Isles secondary production in the benthos is generally high, but shows seasonal variation (Wood, 1987). Generally, secondary production is highest during summer months, when temperatures rise and primary productivity is at its peak. Spring phytoplankton blooms are known to trigger, after a short delay, a corresponding increase in productivity in benthic communities (Faubel et al., 1983). Some of this production is in the form of reproductive products.

    Recruitment processes

    The dominant species in the biotope are polychaetes and bivalves which, tend to have relatively long-lived planktonic larvae. More detailed information concerning recruitment of important characterizing species is given below:
      Lanice conchilega is a polychaete species with separate sexes. During its life of 1-2 years (Beukema et al., 1978), the species initially passes through two larval stages, of which the last one, when it is known as an aulophora larva, lives about 4-6 weeks in the plankton (Kessler, 1963). Kuhl (1972) reported that the larvae of Lanice conchilega are released between April and October. Experimental data and field studies from the Wadden Sea revealed that the existence of 'hard substrate', preferentially tubes of conspecific adults, was a requirement for initial settlement of Lanice conchilega larvae, although single juveniles were also observed to settle on eroded shells of cockles (Cerastoderma edule) and soft-shelled clams (Mya arenaria) (Heuers, 1998; Heuers et al., 1998). Presumably this was the case following the ice winter of 1995/96 which decimated populations of Lanice conchilega on intertidal sand flats of the Dutch Wadden Sea, as settlement of larvae occurred in the absence of adults in the intertidal and shallow sublittoral (Strasser & Pielouth, 2001). Near bottom water velocity and turbulence are thought to be essential factors determining the spatial and temporal distribution of Lanice conchilega. According to Harvey & Bourget (1995) current velocity over a dense Lanice conchilega tube 'lawn' has to reach a value that causes turbulence and facilitates larval settlement. Lower current speeds reduce turbulence and thus the probability of larval settlement is reduced. The current speed at which turbulence developed depended on the density of the tube 'lawn', tube diameter and surface roughness in combination with currents. Grimm (1999) modelled the spatial and temporal distribution of Lanice conchilega with the intention of exploring mechanisms responsible for the large differences in density of the species on a tidal flat of the Dutch Wadden Sea. Field data collected by Brandt et al. (1995) seemed to tentatively confirm the model assumption made by Grimm (1999) that 'the local density of Lanice conchilega is strongly influenced by the overall velocity of the near bottom flow: low density occurs in areas where low velocities prevail, whereas high densities occur where high velocities prevail'. Brandt et al. (1995) recorded a mean flow velocity of 10 cm/s in low density stands of the species, and 20 cm/s near populations of high density.
    • Hayward (1994) summarized recruitment of Arenicola marina. The species breeds late in the year and has a protracted spawning period between September and November, consisting of two peaks, in late September and late November, with a five week period between during which no spawning occurs. Production of eggs starts in February and March, so that by July the entire population consists of sexually mature adults, usually equal in number. Spawning is synchronous, induced by falling seawater temperature at a threshold of 13 °C, or by an abrupt downward temperature shock (Farke & Berghuis, 19790. Males are the first to spawn into their burrows, from which it is ejected to form pools on the sand. Females draw sperm into their burrows as they ventilate their burrows and eggs are fertilized. Development and hatching occurs within the female burrow. After spawning males fast for 2 days while females fast for 3-4 weeks, presumably to avoid ingesting eggs and larvae (Farke & Berghuis, 1979). The larvae leave the burrow when they have grown three chaete-bearing body segments. Post larvae are capable of active migration by crawling, swimming in the water column and passive transport by currents e.g. Günther (1992) suggested that post-larvae of Arenicola marina were transported distances in the range of 1 km. The species may live between 5 and 10 years.
    • Reproductive data concerning Magelona mirabilis is scarce (Fiege et al., 2000). It is generally agreed that Magelona mirabilis displays characteristics typical of an r-selected species, i.e. high reproductive rate, short life span and high dispersal potential (Krönke, 1990; Niermann et al., 1990), and typically occurs in the early successional stages of variable, unstable habitats (Bosselmann, 1989; Niermann et al., 1990). Magelona mirabilis seems to have a protracted reproductive period. Fiege et al. (2000) recorded males with sperm masses and females with eggs in Scotland in March and egg bearing females in France in May. Probert (1981) reported Magelona sp. larvae in Plymouth Sound between April and November with greatest abundances from July to October, and Bosselmann (1989) reported that Magelona sp. larvae were captured in plankton trawls on most dates of sampling in the German Bight but were most abundant in August and September. Bosselmann (1989) also noted large interannual variability in numbers of Magelona sp. larvae in the plankton.
    • The bivalves in the biotope are gonochoristic broadcast spawners with pelagic larval dispersal. They therefore have the potential to recruit both locally and remotely. However, bivalve populations typically show considerable pluriannual variations in recruitment, suggesting that recruitment is patchy and/or post settlement processes are highly variable (e.g. Dauvin, 1985). Abra alba is generally considered to be an 'r-strategist'; capable of rapidly exploiting any new or disturbed substratum suitable for colonization through larval recruitment, secondary settlement of post metamorphosis juveniles or redistribution of adults following storms (Rees & Dare, 1993). Normally, there two distinct spawning periods, in July and September and according to the season of settlement, individuals differ in terms of growth and potential life span (Dauvin & Gentil, 1989). Autumn settled individuals from the Bay of Morlaix, France, initially showed no significant growth; they were not collected on a 1 mm mesh sieve until April, 5 to 7 months after settlement. Such individuals were expected to have a maximum life span of 21 months and could produce two spawnings. In contrast, veliger larvae that settled during the summer grew very rapidly and were collected on a 1 mm mesh sieve just one month after settlement. They lived about one year and spawned only once (Dauvin & Gentil, 1989).
    • Time for community to reach maturity

      The life history characteristics of the species, particularly the polychaetes, which characterize the biotope suggest that the community would probably reach maturity within 3 years. For instance, settlement of Lanice conchilega has been reported to be more successful in areas with existent adults than areas without (Heuers & Jaklin, 1999), however, Strasser & Pielouth (2001) reported settlement in location without adults, but that establishment of a mature population took three years. Adults of Arenicola marina reach sexual maturity by their second year (Newell, 1948; Wilde & Berghuis, 1979) but may mature by the end of their first year in favourable conditions depending on temperature, body size, and hence food availability (Wilde & Berghuis, 1979). The life history characteristics of Abra alba and its widespread distribution contribute to its powers of recoverability. Diaz & Castaneda et al., (1989) experimentally investigated recolonization sequences of benthic areas over a period of one year following defaunation of the sediment. Recovery of the Abra alba population was rapid, recruitment occurred from surrounding populations via the plankton. The abundance, total biomass and diversity of the community all increased until a maximum was reached after 20 to 24 weeks, according to the season. The community within the experimental containers matched that of the surrounding areas qualitatively but quantitatively within 4 to 8 months depending on the seasonal availability of recruits, food supply and faunal interactions. The experimental data suggests that Abra alba would colonize available sediments within the year following environmental perturbation. Summer settled recruits may grow very rapidly and spawn in the autumn, whilst autumn recruits experience delayed growth and may not reach maturity until the following spring/summer.

      Additional information

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    Preferences & Distribution

    Recorded distribution in Britain and IrelandPresent on all coasts of Britain with the exception of the north east coast of Scotland and south east coast of England. Reported in Ireland at Rosbin Cove in Roaringwater Bay, and along the coast from Kilmichael Point southwards around Cahore Point to Raven Point, County Wexford.

    Habitat preferences

    Depth Range
    Water clarity preferences
    Limiting Nutrients Data deficient
    Biological Zone
    Other preferences Tidal streams

    Additional Information

    The dense Lanice biotope (LGS.Lan) on certain lower shores may be a littoral extension of this biotope.

    Species composition

    Species found especially in this biotope

    Rare or scarce species associated with this biotope


    Additional information

    The MNCR recorded ca 613 species within this biotope, although not all species occur in all examples of the biotope (JNCC, 1999). Connor et al (1997b) noted that this biotope may consist of a number of separate communities. Although the density of Lanice conchilega is the characteristic feature of the biotope, community composition probably varies with location, the degree of detritus and nutrient input and the local hydrographic regime, resulting in the high number of species recorded in this biotope.

    Sensitivity reviewHow is sensitivity assessed?


    Lanice conchilega is the dominant polychaete within the biotope. It qualifies as an 'ecosystem engineer' in that it changes and/or creates a habitat, which affects other organisms (Jones et al., 1994; 1997). The tubes of Lanice conchilega, protruding 2-3 cm above the sediment surface, strongly affect the hydrodynamic regime in the benthic boundary layer and thus the distribution of co-occurring biota (see ecological interactions and habitat complexity) (e.g. Eckman et al., 1981; Eckman, 1985; Carey, 1983, 1987; Zühlke et al., 1998; Zühlke, 2001). Loss or reduction of the Lanice conchilega population would probably result in loss of the biotope as described and the species has been assessed to be a key structuring species. The biotope is also characterized by a number of polychaete species, therefore, Arenicola marina has been included as an important characterizing species but reference has also been made to other polychaetes species where possible. The sensitivity of bivalve species has been represented by Abra alba which can be common in this biotope.

    Species indicative of sensitivity

    Community ImportanceSpecies nameCommon Name
    Important otherAbra albaA bivalve mollusc
    Important characterizingArenicola marinaBlow lug
    Key structuralLanice conchilegaSand mason worm

    Physical Pressures

     IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
    High High Moderate Major decline High
    Characterizing species in the biotope are infaunal and would therefore be removed along with the substratum. Some epifaunal and swimming species, such as amphipods and the harbour crab Liocarcinus depurator, may be able to avoid the factor. However, because the species which characterize the biotope would be lost, intolerance has been assessed to be high and there would be a major decline in species richness. Recoverability has been assessed to be high (see additional information below).
    Low Immediate Not sensitive No change Moderate
    The tube of Lanice conchilega rises several centimetres above the sediment surface. Ziegelmeier (1952) showed that the polychaete increased the height of the tube top with increasing sedimentation. It is therefore, unlikely that silt would smother the worm. For other polychaetes, such as Magelona mirabilis, that deposit feeds at the surface by extending contractile palps from its burrow, a layer of sediment would result in a temporary cessation of feeding activity. Abra alba and Fabulina fabula are shallow burrowers in sandy sediments. These bivalves require their inhalant siphons to be above the sediment surface for feeding and respiration. Smothering with 5 cm of sediment would temporarily halt feeding and respiration and require the species to relocate to its preferred depth. Similarly, infaunal polychaete species would move up through additional sediment without adverse effect. Intolerance has been assessed to be low as relocation would be at energetic cost and feeding activity would be inhibited. Recovery has been assessed to be immediate. Smothering by viscous or impenetrable materials would be expected to have a more severe effect.
    Tolerant Not relevant Not relevant Rise Moderate
    Suspension feeding species within the biotope are likely to benefit from an increase in suspended sediment especially if there was a significant proportion of organic matter in the suspended sediment and if food was previously limiting. Lanice conchilega uses its 'feeding crown' to attain particles and unless the 'feeding crown' becomes clogged and requires excessive cleaning at energetic cost the species is unlikely to be adversely affected. Infaunal species such as Arenicola marina are unlikely to be perturbed. An assessment of not sensitive * has been made as suspension feeders may benefit from the increased availability of food.
    Low High Moderate Decline Low
    A decrease in suspended sediment would reduce the amount of available food for suspension feeders such as Lanice conchilega and bivalve molluscs. Deposit feeders such as Arenicola marina are unlikely to be directly affected, although a reduction in suspended sediment may eventually cause deposits of organic matter to become limiting as a consequence of reduced supply. An intolerance assessment of low has been made as suspension feeding species would experience reduced growth and fecundity rather than mortality over the period of one month. On return to prior conditions optimal feeding is likely to resume and recoverability has been assessed to be very high.
    Low High Low No change Low
    Normally, exposure to direct sunlight and air is unlikely as the biotope is subtidal. However, desiccation would become a stress if sediments in the very shallow sublittoral became 'baked' by sunlight during a prolonged low tide event or the component species in the biotope were to be removed from the sediment and stranded, unable to reburrow. However, the majority of the fauna, polychaete worms and bivalves live infaunally and so are likely to be protected from desiccation stress. Bivalves respond to desiccation stress by valve adduction and it is likely that they would be able to retain enough water within their shells to avoid mortality during the benchmark period of one hour. Mobile species would migrate. In avoiding the effects of desiccation the fauna would not be able to feed and respiration would be compromised, so there is likely to be some energetic cost. Therefore intolerance has been assessed to be low. On immersion, metabolic activity should quickly return to normal and recoverability is therefore recorded as very high. There is unlikely to be a decline in species richness.
    Tolerant Not relevant Not relevant No change Moderate
    The biotope occurs from the shallow sublittoral (0-5 m) down to a depth of 20 m, therefore it does not normally experience periods of emergence. An increase in emergence is unlikely to occur in this biotope, but if it did only the uppermost part of it in the very shallow sublittoral would be affected. However, on certain shores the A2.245 (Dense Lanice conchilega in tide-swept lower shore sand) may be a littoral extension of this biotope. The LGS.Lan biotope comprises of many similar functioning species (if not the same) suggesting that species of the IGS.Lcon biotope would be not sensitive. An increase in emergence may however lead to an increase in predation (of the relatively small proportion of the biotope affected) by wading sea birds. However, on balance the major extent of the biotope would be unaffected and so an assessment of not sensitive has been made.
    Not sensitive* Not relevant
    The biotope occurs from the shallow sublittoral (0-5 m) down to a depth of 20 m where it is continually immersed and therefore would not be affected by a decrease in emergence regime.
    High High Moderate Decline Low
    The nature of the substratum is, in part, determined by the hydrographic regime including water flow rate. Changes in the water flow rate will change the sediment structure and have concomitant effects on the community, as many sediment dwelling species have defined substratum preferences (e.g. Bathyporeia pelagica).
    However, moderate to high velocities of water flow have been reported to enhance settlement of Lanice conchilega larvae (Harvey & Bourget, 1995) (see recruitment processes). But an increase in water flow from e.g. moderately strong to very strong, would probably winnow away smaller particulates, increasing average particle size in favour of gravels and pebbles. Therefore, the density of the Lanice conchilega population may decline, in part due to lack of suitable substrata with which to build its tubes, and partly from interference with its feeding. The community would probably become dominated by water flow tolerant species, that prefer coarse substratum, while species such as Arenicola marina, Abra alba, and Spiophanes bombyx may be excluded. The biotope may start to resemble the burrowing anemone dominated community A5.132. Therefore, an intolerance of high has been recorded. On return to prior conditions recoverability is likely to be high (see additional information below).
    Tolerant Not sensitive* No change Low
    The nature of the substratum is, in part, determined by the hydrographic regime including water flow rate. Changes in the water flow rate will change the sediment structure and have concomitant effects on the community.

    Reduced water flow is a factor that has been identified as affecting the density of Lanice conchilega. Recruitment to the benthos is reduced under low flow as a result of reduced turbulence (Harvey & Bourget, 1995) (see recruitment processes). Furthermore, at the benchmark level, decreased water flow rate would probably increase deposition of finer sediments, and increase siltation. The sediment would probably begin to favour deposit feeders and detritivores, to the detriment of the suspension feeders. The average grain size of the sediment would be reduced, and the community may start to be replaced over a period of one year by communities characteristic of muddy sands, with a higher proportion of deposit feeding species, perhaps e.g. A5.331 or A5.241. Therefore, an intolerance of high has been recorded. On return to prior conditions recoverability has been assessed to be high (see additional information below)

    Low Very high Very Low No change Low
    The geographic distribution of polychaete and bivalve species characteristic of this biotope extend to the south of the British Isles, so are likely to be tolerant of a long-term chronic temperature increase of 2 °C. Infaunal species are likely to be protected to some extent from direct effects of acute increases in temperature, although increased temperatures may affect infauna indirectly, by stimulating increased bacterial activity and increased oxygen consumption. Depletion of oxygen from the interstitial waters would result in reduced oxygen levels (hypoxia) or more severely the complete absence of oxygen (anoxia) (see deoxygenation) in the sediment (Hayward, 1994).
    Lethal temperatures (LT50) have been reported for species such as Abra alba and Fabulina fabula (see MarLIN reviews) but temperatures in excess of 20 °C are not likely around the British Isles (Hiscock, 1998). An acute increase in temperature at the benchmark level may result in physiological stress endured by the species but is unlikely to lead to mortality. The biotope is subtidal and probably protected from extremes of temperature by the depth of overlying water.
    Therefore, an intolerance of low has been recorded to represent sub-lethal effects on growth and reproduction. A recoverability of very high has been suggested.
    Low Very high Moderate Decline Low
    Lanice conchilega is intolerant of low temperatures (Beukema, 1990). An intertidal population of Lanice conchilega, in the northern Wadden Sea, was wiped out during the severe ice winter of 1995/96 (Strasser & Pielouth, 2001). The population took three years to fully recover, as there was low recruitment for the first two years. Crisp (1964) described mortality of Lanice conchilega between the tidemarks but not at lower levels during the severe winter of 1962/63.
    Other characterizing species in the biotope are recorded north of the British Isles (e.g. Arenicola marina, Abra alba, and Spiophanes bombyx) and are unlikely to be affected by long term chronic decreases in temperature. However, Arenicola marina may be more intolerant, it experienced some mortality at 5 °C in laboratory studies, although in the field it would derive protection from its infaunal habit.
    Overall, species within the biotope are probably protected from extremes of temperature by their infaunal habit and depth of overlying water. While some more intolerant species may be reduced in abundance or migrate to deeper water, reducing species richness, an overall intolerance of low has been recorded. Recoverability is probably very high.
    Low Very high Very Low No change Low
    Production within the biotope is predominantly secondary, derived from detritus and to some extent phytoplanktonic production. Characteristic infauna do not require light and therefore the effects of increased turbidity on light attenuation are not directly relevant. However an increase in turbidity may affect primary production in the water column and therefore reduce the availability of phytoplankton as food but, phytoplankton would also be transported in to the biotope from distant areas, so the effect of increased turbidity may be mitigated. The increased turbidity persists for a year, so decreased food availability would probably affect growth and fecundity of species and an intolerance of low has been recorded. As soon as light levels return to normal, phytoplanktonic primary production would increase, the species would resume optimal feeding, so recoverability has been assessed to be very high.
    Tolerant* Not sensitive No change Moderate
    It is possible that decreased turbidity would increase primary production in the water column by phytoplankton and by the microphytobenthos. The resultant increase in food availability may enhance growth and reproduction of both suspension and deposit feeding species but only if food was previously limiting. An intolerance assessment of not sensitive* has been made.
    High High Moderate Major decline Moderate
    The biotope occurs in 'sheltered', 'very sheltered' and 'extremely sheltered' locations (Connor et al., 1997a). An increase in wave exposure is likely to have adverse effect on the biotope. Rees et al. (1977) found that only 1 % of the Lanice conchilega population in Colwyn Bay apparently survived after winter storms. Presumably the oscillatory action on the prominent tube served to dislodge the species. An increase in wave exposure would also lead to erosion of the substratum in the shallowest locations, which will alter the extent of suitable habitat available for the community. Intolerance has been assessed to be high as important characterizing species would be lost and the habitat damaged. On return to prior conditions recoverability is likely to be high (see additional information below).
    Tolerant Not sensitive* No change Low
    The biotope occurs in 'sheltered', 'very sheltered' and 'extremely sheltered' locations (Connor et al., 1997a). A further decrease in wave exposure may result in increased siltation and a consequent change in sediment characteristics (Hiscock, 1983). A substratum with a higher proportion of fine sediment would probably result in the increased abundance of the deposit feeders within the biotope, particularly species which favour finer sediments, such as the polychaete Aphelochaeta marioni and the echinoid Echinocardium cordatum. However, in the absence of wave action, tidal flow is likely to be a more significant factor structuring the community, replenishing oxygen, supplying planktonic recruits and would maintain a supply of suspended organic matter in suspension for suspension feeders. Therefore the biotope has been assessed to be not sensitive.
    Tolerant Not relevant Not relevant No change Low
    No information was found concerning the intolerance of the biotope or the characterizing species to noise. The siphons of bivalves and palps of polychaetes are likely to detect vibrations and are probably withdrawn as a predator avoidance mechanism. However, it is unlikely that the biotope will be affected by noise or vibrations caused by noise at the level of the benchmark.
    Tolerant Not relevant Not relevant No change Low
    The majority of species in the biotope are infaunal and have little or no visual acuity. No evidence was found concerning intolerance to visual presence, but it is unlikely that the biotope would be affected.
    Intermediate High Low Minor decline Moderate
    Lanice conchilega inhabits a permanent tube and is likely to be damaged by any activity that penetrates the sediment. Ferns et al. (2000) investigated the effect of mechanical cockle harvesting. The tubes of Lanice conchilega were damaged but this damage was seen to be repaired. In the intertidal, mechanical cockle harvesting resulted in significant losses of common invertebrates in muddy sand and clean sand in the Burry Inlet (Ferns et al. , 2000). For example, losses varied from 31% of Scoloplos armiger to 83% of Pygospio elegans in dense populations. Populations of Nephtys hombergii and Scoloplos armiger took over 50 days to recover. However, recovery was more rapid in clean sand than in muddy sand. In muddy sand, Bathyporeia pilosa took 111 days to recover while Pygospio elegans and Hydrobia ulvae had not recovered their original abundance after 174 days (Ferns et al. , 2000). In a similar study, Hall & Harding (1997) found that non-target benthic fauna recovered within 56 days after mechanised cockle harvesting. However, Hall & Harding (1997) study took place in summer while Ferns et al. (2000) study occurred in winter.

    Despite their apparent robust body form, bivalves are also vulnerable to physical abrasion. For example, as a result of dredging activity, mortality and shell damage has been reported in Mya arenaria and Cerastoderma edule (Cotter et al. , 1997). The most sensitive species identified was Echinocardium cordatum which has a fragile test that is likely to be damaged by an abrasive force such as movement of trawling gear over the seabed. A substantial reduction in the numbers of Echinocardium cordatum due to physical damage from scallop dredging has been observed (Eleftheriou & Robertson, 1992). The species has high fecundity, normally reproduces every year and has pelagic larvae so recovery would be expected. Intolerance has been assessed to be intermediate, as some mortality would be expected as a result of abrasion and physical disturbance. Recoverability has been assessed to be high (see additional information below).

    Intermediate Immediate Very Low Minor decline Moderate
    Yonow (1989) observed Lanice conchilega re-establishing tubes immediately after removal from the sediment, when placed on a suitable sediment in the laboratory. Abra alba, Fabulina fabula and Magelona mirabilis are all active burrowers and are capable of reburying themselves if displaced to the surface of a suitable substratum (Jones, 1968; Salzwedel, 1979). However, while at the sediment surface they are vulnerable to predation from crabs, echinoderms (Aberkali & Trueman, 1985) and bottom feeding fish (Hunt, 1925; Hayward & Ryland, 1995) so there is likely to be some mortality. Intolerance has been assessed to be intermediate. However, it is likely that the majority of displaced specimens would obtain protection within the substratum relatively quickly so recoverability has been assessed to be immediate.

    Chemical Pressures

    High High Moderate Major decline Low
    No evidence of the effects of chemical contaminant on Lanice conchilega were found. However, exposure of Hediste diversicolor and Arenicola marina to Ivermecten resulted in significant mortality (see MarLIN reviews; Collier & Pinn, 1998). Beaumont et al. (1989) investigated the effects of tri-butyl tin (TBT) on benthic organisms. At concentrations of 1-3 µg/l there was no significant effect on the abundance of Hediste diversicolor or Cirratulus cirratus after 9 weeks in a microcosm. However, no juvenile polychaetes were retrieved from the substratum suggesting that TBT had an effect on the larval and/or juvenile stages of these polychaetes. Bryan & Gibbs (1991) reported that Arenicola costata larvae were unaffected by 168 hr exposure to 2000 ng TBT/ l seawater and were probably relatively tolerant, however in another study, Scoloplos armiger exhibited a dose related decline in numbers when exposed to TBT paint particles in the sediment.

    (Møhlenberg & Kiørboe, 1983) demonstrated that pesticide contamination impaired or prevented burrowing in Abra alba, which would probably result in the species being exposed to predatory starfish and fish. Beaumont et al. (1989) concluded that bivalves are particularly intolerant of tri-butyl tin (TBT). For example, when exposed to 1-3 µg TBT/l, Cerastoderma edule and Scrobicularia plana suffered 100 % mortality after 2 weeks and 10 weeks respectively. There is also evidence that TBT causes recruitment failure in bivalves, either due to reproductive failure or larval mortality (Bryan & Gibbs, 1991).

    Pesticides and herbicides were suggested to be very toxic for invertebrates, especially crustaceans (amphipods, isopods, mysids, shrimp and crabs) and fish (Cole et al., 1999). Cole et al. (1999) suggested that TBT was very toxic to algae (including microalgae), molluscs, crustaceans and fish, with observable endocrine disrupting effects in gastropods. Waldock et al. (1999) examined recovery of benthic infauna of the Crouch estuary after a ban on the use of TBT on small boats. They observed marked increase in species diversity, especially of Ampeliscid amphipods and polychaetes (e.g. Tubificoides species and Aphelochaeta marioni) which mirrored the decline in sediment TBT concentration. Whilst a causal link could not be shown, the study by Waldock et al. (1999) suggested that crustacean and polychaete diversity may be inhibited by TBT contamination.

    Polychaete species vary greatly in their tolerance of chemical contamination. However, evidence suggests that the polychaetes within this biotope, including the dominant species Lanice conchilega, are potentially highly intolerance of chemical contamination from pesticides or TBT. The abundance and reproduction of bivalves, crustaceans and other species in the biotope may also be adversely affected. Therefore, an intolerance of high has been recorded, albeit at low confidence. Species richness is likely to decline markedly, due to the dominance of fewer tolerant species. On return to prior conditions and assuming deterioration of the contaminants recoverability is likely to be high (see additional information below).
    Heavy metal contamination
    Intermediate High Low Decline Moderate

    Bryan (1984) suggested that polychaetes are fairly resistant to heavy metals based on the species studied. Short term toxicity in polychaetes was highest to Hg, Cu and Ag, declined with Al, Cr, Zn and Pb whereas Cd, Ni, Co and Se were the least toxic. However, polychaete species vary in their tolerance to heavy metals. For example, exposure to 10 ppm Cd in seawater halted feeding in Arenicola marina but continued at 1 ppm (Rasmussen et al., 1998), while median lethal concentrations (LC50) of 20 µg Cu/g, 50 µg Zn/g, and 25 µg Cd/g have been reported (Bat & Raffaelli, 1998). Arenicola marina was also found to accumulate As, Cd, Sb, Cu, and Cr when exposed to pulverised fuel ash (PFA) in sediments (Jenner & Bowmer, 1990). The spionid polychaete, Aphelochaeta marioni, is apparently very tolerant of heavy metal contamination, occurring in sediments with very high concentrations of arsenic, copper, tin, silver and zinc (Bryan & Gibbs, 1983) and accumulating remarkable concentrations of arsenic (Gibbs et al., 1983). Hediste diversicolor has been found successfully living in estuarine sediments contaminated with copper ranging from 20 µm Cu/g in low copper areas to >4000 µm Cu/g where mining pollution is encountered e.g. Restronguet Creek, Fal Estuary, Cornwall (Bryan & Hummerstone, 1971).

    Bryan (1984) stated that Hg was the most toxic metal to bivalve molluscs while Cu, Cd and Zn seem to be most problematic in the field. In bivalve molluscs, Hg was reported to have the highest toxicity, mortalities occurring above 0.1-1 µg/l after 4-14 days exposure (Crompton, 1997), toxicity decreasing from Hg > Cu and Cd > Zn > Pb and As > Cr ( in bivalve larvae, Hg and Cu > Zn > Cd, Pb, As, and Ni > to Cr). However, bivalves vary in their tolerance to heavy metals.

    Cole et al. (1999) suggested that Hg, Pb, Cr, Zn, Cu, Ni, and Ar were very toxic to invertebrates. Crustaceans are generally regarded to be intolerant of cadmium (McLusky et al., 1986). In laboratory investigations Hong & Reish (1987) observed 96 hour LC50 (the concentration which produces 50 % mortality) of between 0.19 and 1.83 mg/l in the water column for several species of amphipod.

    Overall, polychaetes and bivalves vary in intolerance but may exhibit at least intermediate intolerance to some heavy metals, especially Hg. Amphipods are probably more intolerant, and heavy metal contamination is likely to result in a decline in species richness. On return to prior conditions, and assuming deterioration of the contaminants ,recoverability has been assessed to be high (see additional information below).
    Hydrocarbon contamination
    Intermediate High Low Decline High
    Suchanek (1993) reviewed the effects of oil spills on marine invertebrates and concluded that, in general, on soft sediment habitats, infaunal polychaetes, bivalves and amphipods were particularly affected. A 20 year study investigating community effects after the Amoco Cadiz oil spill of 1978 (Dauvin, 2000) found that a population of Lanice conchilega was established between 1978-84 but disappeared after 1985. Hailey (1995) cited substantial kills of Hediste diversicolor, Cerastoderma edule, Macoma balthica, Arenicola marina and Hydrobia ulvae as a result of the Sivand oil spill in the Humber estuary in 1983.

    Levell (1976) examined the effects of experimental spills of crude oil and oil: dispersant (BP1100X) mixtures on Arenicola marina. Single spills caused 25-50 % reduction in abundance and additional reduction in feeding activity. Up to four repeated spillages (over a 10 month period) resulted in complete eradication of the affected population either due to death or migration out of the sediment. Levell (1976) also noted that recolonization was inhibited but not prevented. Prouse & Gordon (1976) found that Arenicola marina was driven out of the sediment by waterborne concentration of >1 mg/l of fuel oil or sediment concentration of >100 µg/g fuel oil. Seawater oil concentrations of 0.7 mg oil /l reduced feeding after five hours and all worms exposed for 22 hours to 5mg/l oil left the sediment and died after three days. However, the sample size, in the experiment, was very small (6 worms). Sediment concentration >10g/g could reduce feeding. However, Nephtys hombergii, cirratulids and capitellids were largely unaffected by the Amoco Cadiz oil spill Conan (1982).

    Generally, contact with oil in bivalves causes an increase in energy expenditure and a decrease in feeding rate, resulting in less energy available for growth and reproduction. Sublethal concentrations of hydrocarbons also reduce infaunal burrowing rates. After the Amoco Cadiz oil spill Fabulina fabula (studied as Tellina fabula) started to disappear from the intertidal zone a few months after the spill and from then on was restricted to subtidal levels. In the following two years, recruitment of Fabulina fabula was very much reduced (Conan, 1982).

    The Amoco Cadiz oil spill also resulted in reductions in abundance, biomass and production of the community through the disappearance of the dominant populations of the amphipods Ampelisca sp. which are very sensitive to oil contamination (Dauvin, 1998) The sediment rapidly de-polluted and, in 1981, benthic recruitment occurred under normal conditions (Dauvin, 1998). However, the recovery of Ampelisca populations took up to 15 years. This was probably due to the amphipods' low fecundity, lack of pelagic larvae and the absence of local unperturbed source populations (Poggiale & Dauvin, 2001).

    The above evidence suggests that soft sediment communities are highly intolerant of perturbation by oil spills. However, the biotope occurs subtidally and so the majority of the biotope is unlikely to be affected directly but may be exposed to water soluble fractions of hydrocarbons, and oils adsorbed onto particulates. Therefore, an intolerance of intermediate has been recorded. Recovery of amphipods to the biotope is likely to be slow. However, Lanice conchilega was shown to be relatively opportunistic after the Amoco Cadiz oil spill, colonizing shortly after the spill (Dauvin, 2000). Therefore, recoverability is likely to be high.
    Radionuclide contamination
    No information Not relevant No information Not relevant Not relevant
    Changes in nutrient levels
    High High Moderate Decline Low
    Nutrient enrichment can lead to significant shifts in community composition in sedimentary habitats. Typically the community moves towards one dominated by deposit feeders and detritivores, such as polychaete worms (see review by Pearson & Rosenberg, 1978). The IGS.Lcon biotope includes some species tolerant of nutrient enrichment, such as the polychaete Capitella capitata (Pearson & Rosenberg, 1978). It is likely that such species would increase in abundance following nutrient enrichment (Elliott, 1994), with an associated decline in suspension feeding species, e.g. Lanice conchilega and organisms adapted to low nutrient levels, such as Magelona mirabilis (Niermann, 1996). In a sewage dumping region of the North Sea, a great increase in the abundance of Abra alba occurred in much of the dumping area because of the ecological adaptations of the species enabled it to exploit the greatly increased supply of nutrients (Caspers, 1981). In extreme cases of eutrophication, however, sediments may become anoxic and defaunated (see deoxygenation below; Elliott, 1994).
    Indirect effects may include algal blooms. Algal blooms may smother the sediment and result in localised hypoxia and anoxia as a consequence of decomposition and mineralization of organic matter. Algal blooms have been implicated in mass mortalities of Arenicola marina, e.g. in South Wales where up to 99 % mortality was reported (Holt et al. 1995; Olive & Cadman, 1990; Boalch, 1979). The dinoflagellate bloom on the southwest coast of England in summer 1978 was also reported to cause mortalities in bivalves (e.g.Ensis species,) and the heart urchin Echinocardium cordatum (Forster, 1979). Overall, the structure of the community is likely to change in favour of deposit feeders, with an increase in the abundance of opportunistic species and a decrease in species richness. The dense Lanice conchilega bed is likely to be lost and an intolerance of high has been recorded. On return to prior conditions recoverability is likely to be high (see additional information below).
    Not relevant Not relevant Not relevant Not relevant Not relevant
    IGS.Lcon occurs in full salinity conditions (Connor et al., 1997a) and therefore, a further increase in salinity is unlikely. No information was found concerning the intolerance of the characterizing species to hypersaline conditions.
    High High Intermediate Decline Low
    The biotope has only been recorded in full salinity conditions (Connor et al., 1997b). Therefore, a decrease in salinity is likely to result in changes in the community. For example, although Lanice conchilega may be found in estuaries in reduced salinities its occurs at low abundance. Arenicola marina is unable to tolerate salinities below 24 psu and is excluded from areas influenced by freshwater runoff or input (e.g. the head end of estuaries) where it is replaced by Hediste diversicolor (Hayward, 1994). Abra alba is only recorded in full salinity conditions and may be intolerant of reduced salinity. Overall, a change in salinity from e.g. full to reduced (see benchmark) is likely to reduce the characteristic density of Lanice conchilega, result in loss of intolerant species and favour an increase in the abundance of species tolerant of reduced salinities e.g. Nephtys cirrosa, Scoloplos armiger, and Spiophanes bombyx. The biotope may come to resemble the reduced to low salinity biotope A5.222. Therefore, the biotope described may be lost and an intolerance of high has been recorded. On return to prior conditions recoverability is likely to be high (see additional information below).
    Intermediate High Low Decline Moderate
    Nierman et al. (1990) reported changes in a fine sand community for the German Bight in an area with regular seasonal hypoxia. In 1983, oxygen levels were exceptionally low <3mg O2/l in large areas and < 1mg O/l in some areas. Species richness decreased by 30-50 % and overall biomass fell. Hypoxia is likely to result in an increase in the abundance of tolerant species. For example, Niermann et al. (1990) reported that Spiophanes bombyx was found in numbers at some, but not all areas, during the period of hypoxia. Arenicola marina is also tolerant of low oxygen concentrations and even short periods of anoxia (Dales, 1958; Hayward, 1994; Zebe & Schiedek, 1996). Abra alba was found to survive, with a decreased growth rate, exposure to 2.4 -3.5 mg O2,/ l over a 93 day experimental period (Hylland et al., 1996).

    intolerance has been assessed to be intermediate as some individuals in the biotope may perish but many are likely to survive at the benchmark level. Recoverability is likely to be high (see additional information below).

    Biological Pressures

    No information Not relevant No information Not relevant Not relevant
    information was found concerning microbial pathogens and parasites of polychaete species. However, more than 20 viruses have been described for marine bivalves (Sinderman, 1990). Bacterial diseases are more significant in the larval stages and protozoans are the most common cause of epizootic outbreaks that may result in mass mortalities of bivalve populations. Parasitic worms, trematodes, cestodes and nematodes can reduce growth and fecundity within bivalves and may in some instances cause death (Dame, 1996). Data concerning effects on community composition was not found.
    No information Not relevant No information Not relevant Not relevant
    No evidence was found to suggest that important characterizing species of this biotope are threatened by alien species.
    Intermediate Very high Low Decline Moderate
    Commercially exploited species Ensis spp. and Cerastoderma edule occur in this biotope, but at lower densities and are not considered to be characterizing species. Shellfish of marketable size can be harvested both in the intertidal and subtidal more rapidly and efficiently using mechanical methods such as tractor-powered harvesters and suction dredgers than by traditional methods. Hydraulic suction dredgers operate by fluidising the sand using water jets and then lifting the sediment and infauna into a revolving drum for sorting. The tractor-towed dredge utilises a blade between 70 -100 cm wide that penetrates to a depth of between 20-40 cm. Sediment is sorted through a rotating drum cage (Hall & Harding, 1997). Such machinery adversely impacts on non-target infaunal species as they are sucked or displaced from the sediment and sustain damage as 'by-catch'. For instance, Ferns et al. (2000) recorded significant losses of common infaunal polychaetes from areas of intertidal muddy sand sediment worked with a tractor-towed cockle harvester: 31% of the polychaete Scoloplos armiger (initial density of 120 m²) and 83% of Pygospio elegans (initial density 1850 m²) were removed and bird feeding activity increased on harvested areas as gulls and waders took advantage of invertebrates made available. The intolerance of the biological community to this factor has been assessed to be intermediate as mortalities would occur. In the study by Ferns et al. (2000) the population of Pygospio elegans remained depleted for more than 100 days after harvesting, whilst those of Nephtys hombergi, Scoloplos armiger and Bathyporeia spp. were depleted for over 50 days. However, invertebrate populations in clean sand with relatively few Cerastoderma edule, but with more tube-dwelling species such as Lanice conchilega, recovered more quickly. Recoverability has been assessed to be very high.
    High Very high Low Major decline Moderate

    Additional information

    The life history characteristics of the polychaete and bivalve species that characterize the biotope suggest that the biotope would recover from major perturbations within five years. For instance;
    • Lanice conchilega spends up to 60 days in the plankton and could disperse over a wide area. Heuers & Jaklin (1999) found that areas with adult worms or artificial tubes were settled and areas without these structures were not. Strasser & Pielouth (2001) reported that larvae were seen to settle in areas where there were no adults but took 3 years to re-establish the population. Recoverability is, therefore, probably quicker in areas that already have a population of Lanice conchilega but would occur in suitable substratum within only a few years even in the absence of existing populations.
    • Abra alba demonstrates a considerable capacity for recovery. Abra alba spawns at least twice a year over a protracted breeding period, during which time an average sized animal of 11 mm can produce between 15, 000 and 17, 000 eggs. Such egg production ensures successful replacement of the population, despite high larval mortality which is characteristic of planktonic development. Timing of spawning and settlement suggests that the larval planktonic phase lasts at least a month (Dauvin & Gentil, 1989), in which time the larvae may be transported over a considerable distance. Whilst some larvae may settle back into the parent population, the planktonic presettlement period is important for dispersal of the species and spatial separation from the adults also reduces the chances of adult induced mortality on the larvae through adult filter feeding (Dame, 1996). In addition to dispersal via the plankton, dispersal of post-settlement juveniles may occur via byssus drifting (Sigurdsson et al., 1976, see adult distribution) and probably bedload transport (Emerson & Grant, 1991). Niermann et al. (1990) studied the recovery of a fine sand Fabulina fabula community from the German Bight following a severe hypoxia event. Re-establishment of faunal composition took approximately 8 months, but biomass did not fully recover for approximately 2 years.

    Importance review


    Habitats of Principal ImportanceSubtidal sands and gravels
    Habitats of Conservation ImportanceSubtidal sands and gravels
    UK Biodiversity Action Plan PrioritySubtidal sands and gravels


    None of the species within this biotope are likely to be subject to exploitation in this habitat.

    Additional information



    1. Aberkali, H.B. & Trueman, E.R., 1985. Effects of environmental stress on marine bivalve molluscs. Advances in Marine Biology, 22, 101-198.
    2. Ansell, A.D., 1995. Surface activity of some benthic invertebrate prey in relation to the foraging activity of juvenile flatfishes. In Proceedings of the 28th European Marine Biology Symposium, Institute of Marine Biology, Crete. 23-28 September 1993. Biology and Ecology of Shallow Coastal Waters (ed. A. Eleftheriou, A.D. Ansell & C.J. Smith), pp. 245-252. Fredensborg: Olsen & Olsen
    3. Bandeira, S.O., 1995. Marine botanical communities in southern Mozambique: Sea grass and seaweed diversity and conservation. Ambio, 24, 506-509.
    4. Barnes, R.S.K. & Hughes, R.N., 1992. An introduction to marine ecology. Oxford: Blackwell Scientific Publications.
    5. Bat, L. & Raffaelli, D., 1998. Sediment toxicity testing: a bioassay approach using the amphipod Corophium volutator and the polychaete Arenicola marina. Journal of Experimental Marine Biology and Ecology, 226, 217-239.
    6. Beaumont, A.R., Newman, P.B., Mills, D.K., Waldock, M.J., Miller, D. & Waite, M.E., 1989. Sandy-substrate microcosm studies on tributyl tin (TBT) toxicity to marine organisms. Scientia Marina, 53, 737-743.
    7. Beukema, J.J., 1990. Expected effects of changes in winter temperatures on benthic animals living in soft sediments in coastal North Sea areas. In Expected effects of climatic change on marine coastal ecosystems (ed. J.J. Beukema, W.J. Wolff & J.J.W.M. Brouns), pp. 83-92. Dordrecht: Kluwer Academic Publ.
    8. Beukema, J.J., De Bruin, W. & Jansen, J.J.M., 1978. Biomass and species richness of the macrobenthic animals living on the tidal flats of the Dutch Wadden Sea: Long-term changes during a period of mild winters. Netherlands Journal of Sea Research, 12, 58-77.
    9. Boalch, G.T., 1979. The dinoflagellate bloom on the coast of south-west England, August to September 1978. Journal of the Marine Biological Association of the United Kingdom, 59, 515-517.
    10. Bosselmann, A., 1989. Larval plankton and recruitment of macrofauna in a subtidal area in the German Bight. In Reproduction, Genetics and Distributions of Marine Organisms (ed. J.S. Ryland & P.A. Tyler), pp. 43-54.
    11. Brandt, G., Fleßner, J., Glaser, D. et al., 1995. Dokumentation zur hydrographischen Frühjahrs-Meßkampagne 1994 der ökosystemforschung Niedersächsisches Wattenmeer im Einzugsgebeit der otzmer Balje. Hyrographie Nr. 8, Nieders. Landesamt für ökologie - Forschungstelle Küste, Norderney.
    12. Bryan, G.W. & Gibbs, P.E., 1991. Impact of low concentrations of tributyltin (TBT) on marine organisms: a review. In: Metal ecotoxicology: concepts and applications (ed. M.C. Newman & A.W. McIntosh), pp. 323-361. Boston: Lewis Publishers Inc.
    13. Bryan, G.W. & Hummerstone, L.G., 1971. Adaptation of the polychaete Nereis diversicolor to estuarine sediments containing high concentrations of heavy metals. I. General observations and adaption to copper. Journal of the Marine Biological Association of the United Kingdom, 51, 845-863.
    14. 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.
    15. Buchanan, J.B. & Moore, J.B., 1986. A broad review of variability and persistence in the Northumberland benthic fauna - 1971-85. Journal of the Marine Biological Association of the United Kingdom, 66, 641-657.
    16. Buhr, K-J., 1976. Suspension-feeding and assimilation efficiency in Lanice conchilega. Marine Biology, 38, 373-383.
    17. Buhr, K.J. & Winter, J.E., 1977. Distribution and maintenance of a Lanice conchilega association in the Weser estuary (FRG), with special reference to the suspension-feeding behaviour of Lanice conchilega. In Proceedings of the Eleventh European Symposium of Marine Biology, University College, Galway, 5-11 October 1976. Biology of Benthic Organisms (ed. B.F. Keegan, P.O. Ceidigh & P.J.S. Boaden), pp. 101-113. Oxford: Pergamon Press.
    18. Carey, D.A., 1983. Particle resuspension in the benthic boundary layer induced by flow around polychaete tubes. Canadian Journal of Fisheries and Aquatic Sciences, 40 (Suppl. 1), 301-308.
    19. Carey, D.A., 1987. Sedimentological effects and palaeoecological implications of the tube-building polychaete Lanice conchilega Pallas. Sedimentology, 34, 49-66.
    20. Caspers, H., 1981. Long-term changes in benthic fauna resulting from sewage sludge dumping in the North Sea. Water Science and Technology, 13, 461-479.
    21. Cole, S., Codling, I.D., Parr, W. & Zabel, T., 1999. Guidelines for managing water quality impacts within UK European Marine sites. Natura 2000 report prepared for the UK Marine SACs Project. 441 pp., Swindon: Water Research Council on behalf of EN, SNH, CCW, JNCC, SAMS and EHS. [UK Marine SACs Project.],
    22. Collier, L.M. & Pinn, E.H., 1998. An assessment of the acute impact of the sea lice treatment Ivermectin on a benthic community. Journal of Experimental Marine Biology and Ecology, 230, 131-147.
    23. Conan, G., 1982. The long-term effects of the Amoco Cadiz oil spill. Philosophical Transactions of the Royal Society of London B, 297, 323-333.
    24. 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.
    25. Costa, M.J. & Elliot, M., 1991. Fish usage and feeding in two industrialised estuaries - the Tagus, Portugal and the Forth, Scotland. In Estuaries and Coasts: Spatial and Temporal Intercomparisons (ed. B. Knights & A.J. Phillips), pp. 289-297. Denmark: Olsen & Olsen.
    26. Cotter, A.J.R., Walker, P., Coates, P., Cook, W. & Dare, P.J., 1997. Trial of a tractor dredger for cockles in Burry Inlet, South Wales. ICES Journal of Marine Science, 54, 72-83.
    27. Crompton, T.R., 1997. Toxicants in the aqueous ecosystem. New York: John Wiley & Sons.
    28. Dales, R.P., 1958. Survival of anaerobic periods by two intertidal polychaetes, Arenicola marina (L.) and Owenia fusiformis Delle Chiaje. Journal of the Marine Biological Association of the United Kingdom, 37, 521-529.
    29. Dame, R.F.D., 1996. Ecology of Marine Bivalves: an Ecosystem Approach. New York: CRC Press Inc. [Marine Science Series.]
    30. Dauvin, J-C. & Gentil, F., 1989. Long-term changes in populations of subtidal bivalves (Abra alba and Abra prismatica) from the Bay of Morlaix (Western English Channel). Marine Biology, 103, 63-73.
    31. Dauvin, J.C., 1985. Dynamics and production of a population of Venus ovata (Pennant) (Mollusca-Bivalvia) of Morlaix Bay (western English Channel). Journal of Experimental Marine Biology and Ecology, 91, 109-123.
    32. Dauvin, J.C., 1998. The fine sand Abra alba community of the Bay of Morlaix twenty years after the Amoco Cadiz oil spill. Marine Pollution Bulletin, 36, 669-676.
    33. Dauvin, J.C., 2000. The muddy fine sand Abra alba - Melinna palmata community of the Bay of Morlaix twenty years after the Amoco Cadiz oil spill. Marine Pollution Bulletin, 40, 528-536.
    34. 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.
    35. Dittmann, S., 1999. Biotic interactions in a Lanice conchilega dominated tidal flat. In The Wadden Sea ecosystem, (ed. S. Dittmann), pp.153-162. Germany: Springer-Verlag.
    36. Dyer, K.R., 1998. Estuaries - a Physical Introduction. John Wiley & Son, Chichester.
    37. Eckman, J.E., 1985. Flow perturbation by a protruding animal tube affects sediment bacterial recolonization. Journal of Marine Research, 43, 419-435.
    38. Eckman, J.E., Nowell, A.R.M. & Jumars, P.A., 1981. Sediment destabilization of animal tubes. Journal of Marine Research, 39, 361-374.
    39. Elliot, M., Nedwell, S., Jones, N.V., Read, S.J., Cutts, N.D. & Hemingway, K.L., 1998. Intertidal sand and mudflats & subtidal mobile sandbanks (Vol. II). An overview of dynamic and sensitivity for conservation management of marine SACs. Prepared by the Scottish Association for Marine Science for the UK Marine SACs Project.
    40. Elliott, M., 1994. The analysis of macrobenthic community data. Marine Pollution Bulletin, 28, 62-64.
    41. Emerson, C.W. & Grant, J., 1991. The control of soft-shell clam (Mya arenaria) recruitment on intertidal sandflats by bedload sediment transport. Limnology and Oceanography, 36, 1288-1300.
    42. Everett, R.A., 1991. Intertidal distribution of infauna in a central California lagoon: the role of seasonal blooms of macroalgae. Journal of Experimental Marine Biology and Ecology, 150, 223-247.
    43. Farke, H. & Berghuis, E.M., 1979. Spawning, larval development and migration behaviour of Arenicola marina in the laboratory. Netherlands Journal of Sea Research, 13, 512-528.
    44. Faubel, A., Hartig, E. & Thiel, H., 1983. On the ecology of the benthos of sublittoral sediments, Fladen Ground, North Sea. 1. Meiofauna standing stock and estimation of production. Meteor Forschungsergebnisse, 36, 35-48.
    45. Ferns, P.N., Rostron, D.M. & Siman, H.Y., 2000. Effects of mechanical cockle harvesting on intertidal communities. Journal of Applied Ecology, 37, 464-474.
    46. Fiege, D., Licher, F. & Mackie, A.S.Y., 2000. A partial review of the European Magelonidae (Annelida : Polychaeta) Magelona mirabilis redefined and M. johnstoni sp. nov. distinguished. Journal of the Marine Biological Association of the United Kingdom, 80, 215-234.
    47. Forster, S. & Graf, G., 1995. Impact of irrigation on oxygen flux into the sediment: intermittent pumping by Callianassa subterranea and "piston pumping" by Lanice conchilega. Marine Biology, 123, 335-346.
    48. Grimm, V., 1999. Modelling the spatial and temporal distribution of Lanice conchilega. In The Wadden Sea: stability, properties and mechanisms, (ed. S. Dittmann), pp.147-152. Germany: Springer-Verlag.
    49. Günther, C-P., 1992. Dispersal of intertidal invertebrates: a strategy to react to disturbances of different scales? Netherlands Journal of Sea Research, 30, 45-56.
    50. Hailey, N., 1995. Likely impacts of oil and gas activities on the marine environment and integration of environmental considerations in licensing policy. English Nature Research Report, no 145., Peterborough: English Nature.
    51. Hall, S.J. & Harding, M.J.C., 1997. Physical disturbance and marine benthic communities: the effects of mechanical harvesting of cockles on non-target benthic infauna. Journal of Applied Ecology, 34, 497-517.
    52. Harvey, M. & Bourget, E., 1995. Experimental evidence of passive accumulation of marine bivalve larvae on filamentous epibenthic structures. Limnology and Oceanography, 40, 94-104.
    53. Hayward, P.J. 1994. Animals of sandy shores. Slough, England: The Richmond Publishing Co. Ltd. [Naturalists' Handbook 21.]
    54. Heuers, J. & Jaklin, S., 1999. Initial settlement of Lanice conchilega. Senckenbergiana Maritima, 29 (suppl.), 67-69.
    55. Heuers, J., 1998. Ansiedlung, Dispersion, Rekrutierung und Störungen als strukturierende Faktoren benthischer Gemeinschaften im Eulitoral. Dissertation, Universität Bonn., Dissertation, Universit&aulm;t Bonn.
    56. Heuers, J., Jaklin, S., Zülkhe, R., Dittmann, S., Günther, C-P., Hildenbrandt, H. & Grimm, V., 1998. A model on the distribution and abundance of the tube-building polychaete Lanice conchilega (Pallas, 1766) in the intertidal of the Wadden Sea. Verhandlungen Ges Ökologie, 28, 207-215.
    57. 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.
    58. Hong, J. & Reish, D.J., 1987. Acute toxicity of cadmium to eight species of marine amphipod and isopod crustaceans from southern California. Bulletin of Environmental Contamination and Toxicology, 39, 884-888.
    59. Hunt, J.D., 1925. The food of the bottom fauna of the Plymouth fishing grounds. Journal of the Marine Biological Association of the United Kingdom, 13, 560-599.
    60. Hylland, K., Sköld, M., Gunnarsson, J.S. & Skei, J., 1996. Interactions between eutrophication and contaminants. IV. Effects on sediment-dwelling organisms. Marine Pollution Bulletin, 33, 90-99.
    61. JNCC (Joint Nature Conservation Committee), 1999. Marine Environment Resource Mapping And Information Database (MERMAID): Marine Nature Conservation Review Survey Database. [on-line],
    62. Jones, C.G., Lawton, J.H. & Shackak, M., 1994. Organisms as ecosystem engineers. Oikos, 69, 373-386.
    63. Jones, C.G., Lawton, J.H. & Shackak, M., 1997. Positive and negative effects of organisms as ecosystem engineers. Ecology, 78, 1946-1957.
    64. Jones, M.L., 1968. On the morphology, feeding and behaviour of Magelona sp. Biological Bulletin of the Marine Laboratory, Woods Hole, 134, 272-297.
    65. Jones, S.E. & Jago, C.F., 1993. In situ assessment of modification of sediment properties by burrowing invertebrates. Marine Biology, 115, 133-142.
    66. Kessler, M., 1963. Die Entwicklung von Lanice conchilega (Pallas) mit besonderer Berücksichtigung der Lebensweise. Helgolander Wissenschaftliche Meeresuntersuchungen, 8, 425-476.
    67. Kuhl, H., 1972. Hydrography and biology of the Elbe Estuary. Oceanography and Marine Biology: an Annual Review, 10, 225-309.
    68. Levell, D., 1976. The effect of Kuwait Crude Oil and the Dispersant BP 1100X on the lugworm, Arenicola marina L. In Proceedings of an Institute of Petroleum / Field Studies Council meeting, Aviemore, Scotland, 21-23 April 1975. Marine Ecology and Oil Pollution (ed. J.M. Baker), pp. 131-185. Barking, England: Applied Science Publishers Ltd.
    69. Lopez-Jamar, E., Francesch, O., Dorrio, A.V. & Parra, S., 1995. Long term variation of the infaunal benthos of La Coruna Bay (NW Spain): results from a 12-year study (1982-1993). Scientia Marina, 59(suppl. 1), 49-61.
    70. Møhlenberg, F. and Kiørboe, T. 1983. Burrowing and Avoidance Behaviour in Marine Organisms Exposed to Pesticide Contaminated Sediments. Marine Pollution Bulletin,14, 57 - 60.

    71. McDermott, J.J., 1984. The feeding biology of Nipponnemertes pulcher (Johnston) (Hoplonemertea), with some ecological implications. Ophelia, 23, 1-21.
    72. McLusky, D.S., Bryant, V. & Campbell, R., 1986. The effects of temperature and salinity on the toxicity of heavy metals to marine and estuarine invertebrates. Oceanography and Marine Biology: an Annual Review, 24, 481-520.
    73. NBN (National Biodiversity Network), 2002. National Biodiversity Network gateway., 2008-10-31
    74. Newell, G.E., 1948. A contribution to our knowledge of the life history of Arenicola marina L. Journal of the Marine Biological Association of the United Kingdom, 28, 554-580.
    75. Niermann, U., 1996. Fluctuation and mass occurrence of Phoronis muelleri (Phoronidea) in the south-eastern North Sea during 1983-1988. Senckenbergiana Maritima, 28, 65-79.
    76. Niermann, U., Bauerfeind, E., Hickel, W. & Westernhagen, H.V., 1990. The recovery of benthos following the impact of low oxygen content in the German Bight. Netherlands Journal of Sea Research, 25, 215-226.
    77. Olafsson, E.B. & Persson, L.E., 1986. The interaction between Nereis diversicolor (Muller) and Corophium volutator (Pallas) as a structuring force in a shallow brackish sediment. Journal of Experimental Marine Biology and Ecology, 103, 103-117.
    78. Olive, P.J.W. & Cadman, P.S., 1990. Mass mortalities of the lugworm on the South Wales coast: a consequence of algal bloom? Marine Pollution Bulletin, 21, 542-545.
    79. Pearson, T.H. & Rosenberg, R., 1978. Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. Oceanography and Marine Biology: an Annual Review, 16, 229-311.
    80. Poggiale, J.C. & Dauvin, J.C., 2001. Long term dynamics of three benthic Ampelisca (Crustacea - Amphipoda) populations from the Bay of Morlaix (western English Channel) related to their disappearance after the Amoco Cadiz oil spill. Marine Ecology Progress Series, 214, 201-209.
    81. Probert, P.K., 1981. Changes in the benthic community of china clay waste deposits is Mevagissey Bay following a reduction of discharges. Journal of the Marine Biological Association of the United Kingdom, 61, 789-804.
    82. Prouse, N.J. & Gordon, D.C., 1976. Interactions between the deposit feeding polychaete Arenicola marina and oiled sediment. In Proceedings of a Symposium of the American Institute of Biological Sciences, Arlington, Virginia, 1976. Sources, effects and sinks of hydrocarbons in the aquatic environment, pp. 408-422. USA: American Institute of Biological Sciences.
    83. Rasmussen, A.D., Banta, G.T. & Anderson, O., 1998. Effects of bioturbation by the lugworm Arenicola marina on cadmium uptake and distribution in sandy sediments. Marine Ecology Progress Series, 164, 179-188.
    84. Rees, E.I.S., Nicholaidou, A. & Laskaridou, P., 1977. The effects of storms on the dynamics of shallow water benthic associations. In Proceedings of the 11th European Symposium on Marine Biology, Galway, Ireland, October 5-11, 1976. Biology of Benthic Organisms, (ed. B.F. Keegan, P.O Ceidigh & P.J.S. Boaden), pp. 465-474.
    85. Rees, H.L. & Dare, P.J., 1993. Sources of mortality and associated life-cycle traits of selected benthic species: a review. MAFF Fisheries Research Data Report, no. 33., Lowestoft: MAFF Directorate of Fisheries Research.
    86. Rhoads, D.C. & Young, D.K., 1970. The influence of deposit-feeding organisms on sediment stability and community trophic structure. Journal of Marine Research, 28, 150-178.
    87. Ropert, M. & Dauvin, J-C., 2000. Renewal and accumulation of a Lanice conchilega (Pallas) population in the baie des Veys, western Bay of seine. Oceanologica Acta, 23, 529-546.
    88. Salzwedel, H., 1979. Reproduction, growth, mortality and variations in abundance and biomass of Tellina fabula (Bivalvia) in the German Bight in 1975/1976. Veroffentlichungen des Instituts fur Meeresforschung in Bremerhaven, 18, 111-202.
    89. Schories, D. & Reise, K., 1993. Germination and anchorage of Enteromorpha spp. In sediments of the Wadden Sea. Helgolander Meeresuntersuchungen, 47, 275-285.
    90. Sebens, K.P., 1991. Habitat structure and community dynamics in marine benthic systems. In Habitat structure, (ed. S.S. Bell), pp. 211-234. Chapman & Hall.
    91. Sigurdsson, J.B., Titman, C.W. & Davies, P.A., 1976. The dispersal of young post-larval bivalve molluscs by byssus threads. Nature, 262, 386-387.
    92. Sinderman, C.J., 1990. Principle diseases of marine fish and shellfish, 2nd edition, Volume 2. Diseases of marine shellfish. Academic Press, 521 pp.
    93. Strasser, M. & Pielouth, U., 2001. Recolonization pattern of the polychaete Lanice conchilega on an intertidal sandflat following the severe winter of 1995/96. Helgoland Marine Research, 55, 176-181.
    94. Suchanek, T.H., 1993. Oil impacts on marine invertebrate populations and communities. American Zoologist, 33, 510-523.
    95. Tait, R.V. & Dipper, R.A., 1998. Elements of Marine Ecology. Reed Elsevier.
    96. Thouzeau, G., Jean, F. & Del Amo, Y., 1996. Sedimenting phytoplankton as a major food source for suspension-feeding queen scallops (Aequipecten opercularis L.) off Roscoff (western English Channel) ? Journal of Shellfish Research, 15, 504-505.
    97. Thrush, S.F., 1986. Community structure on the floor of a sea-lough: are large epibenthic predators important? Journal of Experimental Marine Biology and Ecology, 104, 171-183.
    98. Tyler, P.A., 1977. Seasonal variation and ecology of gametogenesis in the genus Ophiura (Ophiuroidea: Echinodermata) from the Bristol Channel. Journal of Experimental Marine Biology and Ecology, 30, 185-197.
    99. Waldock, R., Rees, H.L., Matthiessen, P. & Pendle, M.A., 1999. Surveys of the benthic infauna of the Crouch Estuary (UK) in relation to TBT contamination. Journal of the Marine Biological Association of the United Kingdom, 79, 225 - 232.
    100. Warwick, R.M. & Uncles, R.J., 1980. Distribution of benthic macrofauna associations in the Bristol Channel in relation to tidal stress. Marine Biology Progress Series, 3, 97-103.
    101. Wilde de P.A.W.J. & Berghuis, E.M., 1979. Laboratory experiments on growth of juvenile lugworms, Arenicola marina. Netherlands Journal of Sea Research, 13, 487-502.
    102. Woodin, S.A., 1978. Refuges, disturbance and community structure: a marine soft bottom example. Ecology, 59, 274-284.
    103. Yonow, N., 1989. Feeding observations on Acteon tornatilis (Linnaeus) (Opisthobranchia: Acteonidae). Journal of Molluscan Studies, 55, 97-102.
    104. Zühlke, R., 2001. Polychaete tubes create ephemeral community patterns: Lanice conchilega (Pallas, 1766) associations studied over six years. Journal of Sea Research, 46, 261-272.
    105. Zühlke, R., Blome, D., van Bernem, K.H. & Dittmann, S., 1998. Effects of the tube-building polychaete Lanice conchilega (Pallas) on benthic macrofauna and nematodes in an intertidal sandflat. Senckenbergiana Maritima, 29, 131-138.
    106. Zebe, E. & Schiedek, D., 1996. The lugworm Arenicola marina: a model of physiological adaptation to life in intertidal sediments. Helgoländer Meeresuntersuchungen, 50, 37-68.
    107. Ziegelmeier, E., 1952. Beobachtungen über den Röhrenbauvon Lanice conchilega (Pallas) im Experiment und am natürlichen Standort. Helgolander Wissenschaftliche Meeresuntersuchungen, IV. 107-129.


    This review can be cited as:

    Budd, G.C. 2006. Dense Lanice conchilega and other polychaetes in tide-swept infralittoral sand and mixed gravelly sand. 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:

    Last Updated: 28/06/2006