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

28-06-2006
Researched byRebecca McQuillan & Dr Heidi Tillin 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

Summary

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

Description

Dense beds of Lanice conchilega occur in coarse to medium fine gravelly sand in the shallow sublittoral, where there are strong tidal streams or wave action. Several other species of polychaete also occur as infauna e.g. Spiophanes bombyx,Scoloplos armigerChaetozone setosa and Magelona mirabilis. Lanice beds are found in a wide range of habitats including muddier mixed sediment. The dense Lanice biotope (LGS.Lan) on certain lower shores may be a littoral extension of the current biotope. The presence of L. conchilega in high numbersmay, over time, stabilise the sediment to the extent where a more diverse community may develop (Wood, 1987). Possibly as a result of this, there is a high level of variation with regard the infauna found in SCS.SLan. It is likely that a number of sub-biotopes may subsequently be identified for this biotope. Offshore from the Wash and the North Norfolk coast Lanice beds are often found intermixed with Sabellaria spinulosa beds in muddier mixed sediment, particularly in the channels between the shallow sandbanks, which are so prevalent in this area (IECS, 1995; NRA, 1995). It is possible that the presence of Lanice has stabilised the habitat sufficiently to allow the deposition of finer material, which has subsequently assisted the development of S. spinulosa. It may be more accurate to define SLan as an epibiotic biotope which overlays a variety of infaunal biotopes (e.g. NcirBat in finer sands and AalbNuc or FfabMag in slightly muddier areas) (JNCC, 2015).

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

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

Additional information

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JNCC

Habitat review

Ecology

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, 1977a; 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, 1979). 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

No text entered.

Preferences & Distribution

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.

Habitat preferences

Depth Range 0-5 m, 10-20 m, 5-10 m
Water clarity preferencesField Unresearched
Limiting Nutrients Data deficient
Salinity Full (30-40 psu)
Physiographic
Biological Zone Infralittoral
Substratum Fine clean sand, Gravelly sand, Medium clean sand
Tidal Moderately Strong 1 to 3 knots (0.5-1.5 m/sec.), Strong 3 to 6 knots (1.5-3 m/sec.), Very Weak (negligible), Weak < 1 knot (<0.5 m/sec.)
Wave Exposed, Moderately exposed, Sheltered
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?

Sensitivity characteristics of the habitat and relevant characteristic species

The biotope description and characterizing species are based on JNCC (2015). The biotope is characterized by dense beds of Lanice conchilega occuring in coarse to medium fine gravelly sand in the shallow sublittoral, where there are strong tidal streams or wave action. The sensitivity assessments are based on Lanice conchilega, as the loss or reduction of the Lanice conchilega population would result in a significant change to the character of the biotope and lead to biotope reclassification. The tubes of this species create physical structure in this biotope and modify habitat conditions, this species is therefore considered to be a key structuring/functional species. Several other species of polychaete also occur as infauna e.g. Spiophanes bombyx, Scoloplos armiger, Chaetozone setosa and Magelona mirabilis and the sensitivities of these species are assessed generally.  

Resilience and recovery rates of habitat

Recovery characteristics of Lanice conchilega were assessed previously by Ager (2008) and Callaway et al. (2010). Lanice conchilega is sessile and larval recolonization is, therefore, the most important recovery mechanism. Water transport of adults, intact in tubes, has been observed after storms and this represents a potential colonisation mechanism. The tube itself can be rapidly repaired or rebuilt following damage under variable conditions (Nicolaidou, 2003). Yonow (1989) observed Lanice conchilega re-establishing tubes immediately after removal from the sediment into a suitable sediment in the laboratory.

The larvae of Lanice conchilega spend up to 60 days as plankton and therefore have a wide dispersal potential. Larvae preferentially settle on the tubes of Lanice conchilega but other suitable settlement surfaces might be selected, including artificial tubes (Heuers & Jaklin, 1999) or bivalve shells (Herlyn et al., 2008). Given the preference for settlement on the tubes of adults, recovery of Lanice conchilega populations will be enhanced by the presence of adults that survive impacts.

Population recovery times appear to range between one and four years (Beukema, 1990; Heuers, 1998; Zühlke, 2001; Callaway et al., 2010). Strasser & Pielouth (2001) reported that larvae were observed to settle in areas where there were no adults and the population was subsequently re-established in three years. The length of recovery time was attributed to a decline in the wider meta-population that reduced larval supply (Strasser & Pieloth, 2001). In areas where there are dense reefs with suitable water currents recovery may, therefore, be more rapid due to the greater larval supply, perhaps taking 1-2 years.

Beukema (1990) reported that following removal of entire intertidal populations (following a cold winter), Lanice conchilega populations recovered rapidly (within 1 and 2 years) and total biomass value increased faster as a consequence of generally high successful recruitment in the Wadden Sea. However, Heuers et al. (1998) observed Lanice conchilega re-established abundance within 3 to 4 years in the Spiekeroog area of the Wadden Sea following severe freezing.  In two of three areas disturbed by cultivation of Manila clams, measurable re-colonization took place after one or two years (Callaway et al., 2010 and references therein). The results tally with other descriptions of Lanice conchilega’s re-colonization strategy,

Recovery of some of the associated species may be more rapid than the recovery of dense patches of Lanice conchilega. Species with opportunistic life strategies (small size, rapid maturation and short-life span of 1-2 years), include the characterizing polychaetes Chaetozone setosa and Spiophanes bombyx.The resilience of Chaetozone spp. were reviewed by MES (2010). has a life-span of 1-2 years and reaches sexual maturity in <1 year. There is little information on the fecundity but the eggs are fertilized externally and may have a significant larval dispersal potential. It shows all the characteristics of an opportunistic species with a short life-span and rapid growth rate. Where the environmental conditions are suitable, Chaetozone setosa is likely to recover to be one of the first genera to recover following disturbance (MES, 2010). Scoloplos armiger and Magelona mirabilis are larger and longer-lived and recovery may be longer but occur within the time same scales as Lanice conchilega.

Scoloplos armiger has a life-span of about four years and reaches maturity at two years.  The sexes are separate and as many as 100-5000 eggs of about 0.25 mm are fertilized externally between February-April. The eggs are attached to the seabed in a gelatinous mass and emerge after three weeks and burrow near the site of release. There may be a very short lecithotrophic pelagic phase in subtidal populations but dispersal is very limited. This genus has a low dispersal potential (MES, 2010). Scoloplos armiger is considered to be a species that characterize the end of the transitional phase and the development of final equilibrium communities following impact or disturbance, rather than an opportunistic species that initially colonizes (Newell et al., 1998). 

Magelona spp. are small-medium sized polychaete worms up to 5-20 cm length free-living within sediments. Magelona lives for about 3 years and reaches maturity at 2 years. There is only one reproductive period, which occurs between May-October. The eggs released by the female are fertilized externally and develop into planktotrophic larvae that settle from June-November. The genus has a high dispersal potential and forms dense communities with a relatively rapid growth rate (MES, 2010).

Resilience assessment.  Where the biotope is severely impacted and the entire assemblage is removed, recovery is assessed as ‘Medium’ as re-establishment of a reef may require up to 3 years, based on the observations of Strasser & Pielouth (2001). Where resistance is assessed as ‘Low’ or ‘Medium’, the presence of adults is considered likely to enhance settlement of juveniles and recovery is assessed as ‘High’ (within 2 years). These assessments are considered to equally apply to the associated macrofauna. 

NB: The resilience and the ability to recover from human induced pressures is a combination of the environmental conditions, the frequency (repeated versus a one off event) and the intensity of the disturbance. Recovery of impacted populations will always be mediated by stochastic events and processes acting over different scales including, but not limited to, local habitat conditions, further impacts and processes such as larval-supply and recruitment between populations. Full recovery is defined as the return to the state of the habitat that existed prior to impact. This does not necessarily mean that every component species has returned to its prior condition, abundance or extent, but that the relevant functional components are present and the habitat is structurally and functionally recognizable as the initial habitat of interest. It should be noted that the recovery rates are only indicative of the recovery potential.  

Hydrological Pressures

 ResistanceResilienceSensitivity
High High Not sensitive
Q: High
A: High
C: Medium
Q: High
A: High
C: High
Q: High
A: High
C: Medium

Limited information on the thermal tolerance of Lanice conchilega was found. Lanice conchilega occurs in temperate regions all around UK and Irish shores in the intertidal and shallow subtidal, populations also occur in the Mediterranean, in the Arabian Gulf and the Pacific and along the south east of the North Sea, but the species is absent from Arctic waters (Ropert & Dauvin, 2000; Connor et al., 2004; Degraer et al., 2008). The geographical range suggests that the species is adapted to variable temperature conditions but local populations are likely to be acclimated to prevailing temperatures.

Magelona mirabilis are widespread on the coasts of Britain and Northern Ireland (Richards, 2007) and their distribution extends to north-west Europe and the Mediterranean (Hayward & Ryland, 1995b). Scoloplos armiger is a species complex as is Chaetozone setosa. Both are widely distributed but populations may be sibling species and exhibit different tolerances. Until recently, Chaetozone setosa was considered cosmopolitan with records world-wide, from the intertidal zone to the deep sea. It is now known that there are several species of eyeless Chaetozone spp. in the north-east Atlantic but the worldwide distribution is unclear. Chambers et al. (2007) assessed numerous records of Chaetozone setosa in the north-east Atlantic, and identified habitat preferences Chatezone setosa  was frequently found in habitats where the mean minimum winter bottom temperature is 5-10°C and the summer maximum is >10°C (Chambers et al., 2007)

Sensitivity assessment. Typical surface water temperatures around the UK coast vary seasonally from 4-19°C (Huthnance, 2010). The associated species are considered likely to be tolerant of acute and chronic increases in temperature at the pressure benchmark. Lanice conchilega and the other characterizing polychaetes have a wide geographic range. The lack of evidence for mass mortalities of intertidal populations in very hot summers (compared with reports for low winter temperatures), suggest that the characterizing and associated species are likely to tolerate a chronic or acute increase at the pressure benchmark).

Low High Low
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

Infaunal species such  Lanice conchilega may be protected to some extent for short periods by the ability to burrow deeper within the sediment. Prolonged freezing temperatures may, however, result in mortalities. In the German coastal area during 1978-79 winter, water temperature remained below 0°C on 45 successive days and resulted in the eradication of Lanice conchilega. Up to July 1980 no recolonization of the area had taken place (Buhr, 1981). An intertidal population of Lanice conchilega, in the northern Wadden Sea, was wiped out during the severe winter of 1995/96 (Strasser & Pielouth, 2001), and Crisp (1964) described mortality of Lanice conchilega between the tidemarks but not at lower levels during the severe winter of 1962/63. These severe winters probably exceed the pressure benchmark. Beukema (1990) had noted that the recovery of most species including Lanice conchilega, was rapid (within 1 and 2 years) and total-biomass value increased faster as a consequence of generally high successful recruitment in the Wadden Sea. Whilst Heuers et al. (1998) observed Lanice conchilega re-established abundance within 3 to 4 years in the Spiekeroog area of the Wadden Sea following severe freezing.

Scoloplos armiger is a species complex as is Chaetozone setosa. Both are widely distributed but populations may be sibling species and exhibit different tolerances. Until recently, Chaetozone setosa was considered cosmopolitan with records world-wide, from the intertidal zone to the deep sea. It is now known that there are several species of eyeless Chaetozone in the north-east Atlantic and the worldwide distribution is unclear. Chambers et al. (2007) assessed numerous records of Chaetozone setosa in the north-east Atlantic. The species is frequently found in habitats where the mean minimum winter bottom temperature is 5-10°C and the summer maximum is >10°C.

Sensitivity assessment. Biotope resistance has been assessed to be ‘Low’ as evidence suggests that populations of the key characterizing species Lanice conchilega are likely to be vulnerable to acute decreases in temperature during winter, resilience is assessed as ‘High’ where some adults remain and sensitivity is assessed as ‘Low’.

No evidence (NEv) No Evidence (NEv) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

This biotope occurs in full (30-35 ppt) salinity (JNCC, 2015). At the pressure benchmark, an increase from full to hypersaline (>40 ppt) was considered but no direct evidence was found to assess the sensitivity of this biotope to a long-term increase.

Salinities within tide swept habitats vary due to weather conditions such as rainfall at low tide and evaporation causing hypersaline conditions on hot days. Intertidal shores within estuarine environments can also experience considerable short term changes in salinities. During summer, the Atlantic French coast rapidly increased in salinity and reached hypersaline conditions (48 ppt) in the upper sediment layer within 1.5 hrs (Juneau et al., 2015).  Lanice conchilega as a tube dwelling polychaete may retract deeper into the sediment to avoid unfavourable conditions but some physiological tolerance for short-term increases in salinity is suggested by its presence in the intertidal.

No direct evidence was found to assess sensitivities of the other characterizing polychaete species. Scoloplos armiger was found at low abundances at the discharge point the discharge point of brine effluents at 47-50 psu in the Canary Islands (Riera et al., 2012). However, in the western Baltic Sea Scoloplos armiger abundance was greatest between 12 psu and 17 psu and reduced abundance with increasing salinity was observed (Gogina et al., 2010). As Scoloplos armiger is a species complex and is not a cosmopolitan species there may be differences in tolerances between populations.

Sensitivity assessment. Species within the biotope may tolerate short periods of high salinity, however, prolonged exposure to hypersalinity may lead to changes in species richness, abundance and biomass and loss of characterizing and associated species. Sensitivity to this pressure is not assessed due to lack of evidence.

 

Medium High Low
Q: High
A: High
C: Medium
Q: High
A: Low
C: High
Q: High
A: Low
C: Medium

The subtidal biotope is found in full salinity (JNCC, 2015), but the intertidal biotope LS.LSa.MuSa.Lan is found in both full and variable salinity. It is therefore considered that a decrease in salinity at the pressure benchmark, from full to variable (18-35 ppt), would not result in loss of Lanice conchilega as this falls within the natural habitat distribution.

Scoloplos armiger shows a lower salinity limit of 10.5 psu (Gogina et al., 2010), suggesting the species is tolerant of a decrease from full to reduced salinity and even the low salinity category in the MNCR scale. Magelona mirabilis occurs in the Baltic Sea (Fiege et al., 2000) where salinity is typically lower than in the open ocean. It is likely that some populations of Magelona mirabilis are adapted to reduced salinity habitats, however, no information on the effects of an overall decrease in salinity were found.

Sensitivity assessment. Little empirical evidence was found to assess sensitivity of Lanice conchilega and other characterizing polychaetes to this pressure at the benchmark level and therefore the assessment was made based on the intertidal biotope. As a reduction may lead to some mortalities and a reduction in growth and reproductive success of the key characterizing species, biotope resistance is assessed as ‘Medium’, recovery is assessed as ‘High’ (following restoration of typical conditions) and resilience is ‘High’. Biotope sensitivity is therefore assessed as ‘Low’.

High High Not sensitive
Q: Low
A: NR
C: NR
Q: High
A: High
C: High
Q: Low
A: Low
C: Low

This biotope is found in areas exposed to tidal streams varying from strong (1.5-3 m/s) to negligible (JNCC, 2015).

There is limited evidence to assess changes in water flow rates for >1 year at the benchmark level. Laboratory flume studies have showed that Lanice conchilega feeding rates on diatoms alter according to flow (Denis et al., 2007). Feeding rates were optimal at 0.15 m/s and lower at flow speeds of 0.04 m/s and 0.27 m/s (Denis et al., 2007). Changes in current velocity at the pressure benchmark are relevant to feeding rates, however, impacts are likely to depend on food supply, population density and proportion of inorganic particles.

The motion of the water is the dominant factor in the transportation of sediment, facilitating erosion and redistribution of materials (Hjulström, 1939). This may change the sediment structure and have associated effects on the intertidal community. Lanice conchilega alter between passive and active feeding depending on the tidal regimes, i.e. in still water, they move their tentacles or in high velocities the fringed tentacles are supported. A switch from active to passive feeding behaviour was observed during the experiments between 4 and 8 cm/s (Denis et al., 2007). Moderate to high velocities of water flow have been reported to enhance settlement of Lanice conchilega larvae (Harvey & Bourget, 1997), increasing average particle size in favour of gravels and pebbles.

Reduced water flow is a factor that has been identified as affecting the density of Lanice conchilega, as recruitment to the benthos is reduced (Harvey & Bourget, 1995). The average grain size of the sediment would also be reduced and probably favour deposit feeders and detritivores, to the detriment of the suspension feeders.

Sensitivity assessment.  A change in water flow rate at the pressure benchmark level of 0.1-0.2 m/s is considered to fall within the range of flow speeds experienced by the biotope. Resistance and resilience are assessed as ‘High’ and the biotope considered ‘Not sensitive’ to a change in water flow at the pressure benchmark level.

 

High High Not sensitive
Q: Low
A: NR
C: NR
Q: High
A: High
C: High
Q: Low
A: Low
C: Low

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 may affect the uppermost part of it. However, on certain shores the LS.LSa.MuSa.Lan biotope may be a littoral extension of this biotope. An increase in emergence may 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 biotope resistance is assessed as ‘High’, resilience as ‘High’ and the biotope is considered ‘Not sensitive’.

High High Not sensitive
Q: High
A: Low
C: NR
Q: High
A: High
C: High
Q: Low
A: Low
C: Low

The biotope is recorded from locations that have variable wave exposures (moderately exposed to sheltered) (Connor et al., 2004). The degree of wave exposure influences wave height, as in more exposed areas with a longer fetch waves would be predicted to be higher. 

Sensitivity assessment. As this biotope is found across a range of wave exposed shores, resistance is assessed as ‘High’ and resilience as ‘High’ to a 3 to 5% change in wave height at the pressure benchmark. By default the biotope is considered ‘Not sensitive’.

Chemical Pressures

 ResistanceResilienceSensitivity
Not relevant (NR) Not relevant (NR) Not sensitive
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

'Not sensitive' at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

Contamination at levels greater than the pressure benchmark may adversely affect the biotope. Bryan (1984) reported that short-term toxicity in polychaetes was highest to Hg, Cu and Ag, declined with Al, Cr, Zn and Pb with Cd, Ni, Co and Se being the least toxic. It was recorded that polychaetes have a range of tolerances to heavy metal levels of Cu, Zn, As and Sn being in the order of 1500-3500 µg/g.

Boilly & Richard (1978) stated that the presence of Magelona mirabilis is indicative of sediments which have been contaminated with iron. Studies on a dredge spoil disposal site in the harbours of Boulogne and Dunkerque in France (Bourgain et al., 1988) found higher densities of Magleona mirabilis three months after the dumping of dredge spoil than after five months, when the metal contamination of the sediments was higher. No information regarding the effect of other metals on this species was found.

Not relevant (NR) Not relevant (NR) Not sensitive
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

‘Not sensitive’ at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

Contamination at levels greater than the pressure benchmark may adversely influences the biotope. Suchanek (1993) reviewed the effects of oil spills and concluded that soft sediment polychaetes, bivalves and amphipods were particularly sensitive. Dauvin (2000) noted that 20 years after the Amoco Cadiz oil spill in 1978, Lanice conchilega was re-established between 1978 and 1984 but disappeared after 1985. A similar delayed response was observed by Sanders et al. (1980) and Kingston et al. (1995) as a result of the Florida oil spill and the Braer oil spill (Gómez Gesteira & Dauvin, 2000).

Gray et al. (1990) found that Scoloplos armiger were a dominant species in uncontaminated soft sediments at a case study site adjacent to the Ekofisk oil field but were not present at contaminated sites.

Not relevant (NR) Not relevant (NR) Not sensitive
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

‘Not sensitive’ at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

No evidence concerning specific effects of synthetic chemical contaminants on Lanice conchilega was found. 

Polychaetes vary greatly in their tolerance of chemical contamination. The persistence of these chemical residues is highly dependent on the matrix and ambient environmental conditions. Generally, residues in water are less likely to be a long-term concern because of photodegendation and dilution to below biological significant concentrations. However, TBT has a high binding affinity to sediments and residues incorporated into the sediment tend to persist for longer periods (Austen & McEvoy, 1997; Huntington et al., 2006).

No evidence (NEv) No Evidence (NEv) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No evidence was found

Not relevant (NR) Not relevant (NR) Not sensitive
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

'Not sensitive' at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

Low High Low
Q: High
A: Low
C: NR
Q: High
A: Low
C: High
Q: High
A: Low
C: Low

Niermann 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 (<3 mg O2/l) in large areas and <1 mg O2/l in some areas. Species richness decreased by 30-50% and overall biomass fell. The abundance of Lanice conchilega was significantly reduced during the period of hypoxia (1-3 mg/O2/dm³). However, the period of hypoxia lasted for about a month and, therefore, exceeds the pressure benchmark. Magelona sp. remained abundant during the hypoxic period and decreased slightly in abundance on resumption of normoxia. Spiophanes bombyx was found in small numbers at some, but not all areas, during the period of hypoxia. Once oxygen levels returned to normal Spiophanes bombyx increased in abundance; the evidence suggests that at least some individuals would survive hypoxic conditions (Niermann et al., 1990)

In laboratory conditions, Scoloplos armiger survived low oxygen conditions for 40 hours (Schöttler & Grieshaber, 1988). No evidence was returned by searches on extended exposure to low levels of dissolved oxygen.

Sensitivity assessment. Based on the available evidence biotope resistance is assessed as ‘Low’ (based on Lanice conchilega) and resilience is assessed as 'High' (following restoration of typical habitat conditions). Biotope resistance is therefore assessed as ‘Low’. Due to the lack of evidence for Lanice conchilega, at the pressure benchmark confidence in applicability is assessed as ’Low’.

Not relevant (NR) Not relevant (NR) Not sensitive
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

This pressure relates to increased levels of nitrogen, phosphorus and silicon in the marine environment compared to background concentrations. The benchmark is set at compliance with WFD criteria for good status, based on nitrogen concentration (UKTAG, 2014). 

Sensitivity assessment. 'Not sensitive' at the pressure benchmark that assumes compliance with good status as defined by the WFD.

High High Not sensitive
Q: Low
A: NR
C: NR
Q: High
A: High
C: High
Q: Low
A: Low
C: Low

The polychaete species characterizing this biotope are all deposit feeders that may benefit from an input of organic material and are likely to be resistant to this pressure at the benchmark. Chaetezone setosa were  typical of enriched sites off the coast of Barcelona that were subject to effluents and sludge disposal from treatment plants (Corbera & Cardell, 1995) and other studies suggest the characterizing polychaetes are unlikely to be affected by organic enrichment at the pressure benchmark. Borja et al. (2000) assessed relative sensitivity of Scoloplos armiger as an ABMI Ecological Group II species (indifferent/tolerant to enrichment). Field studies have also identified Scoloplos armiger as a ‘progressive’ species, i.e. one that shows increased abundance under slight organic enrichment (Leppakoski, 1975 cited in Gray, 1979).

Lanice conchilega was categorised by Borja et al. (2000) as AMBI Group II - 'Species indifferent to enrichment, always present in low densities with non-significant variations with time (from initial state, to slight unbalance)'. This assessment was reviewed by Gittenberger & Van Loon (2011) and changed to AMBI Group III - 'Species tolerant to excess organic matter enrichment. These species may occur under normal conditions, but their populations are stimulated by organic enrichment (slight unbalance situations)'. 

Sensitivity assessment. There is little empirical evidence to quantify the effect of organic enrichment deposits of 100 g C/m²/yr on Lanice conchilega. Based on the ranges presented by Cromey et al. (1998), this benchmark pressure is unlikely to have an influence upon Lanice conchilega and associated species. Biotope resistance is assessed as ‘High’ and resilience as ‘High’ (by default), and the biotope is categorized as ‘Not sensitive’.

Physical Pressures

 ResistanceResilienceSensitivity
None Very Low High
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

All marine habitats and benthic species are considered to have a resistance of ‘None’ to this pressure and to be unable to recover from a permanent loss of habitat (resilience is ‘Very Low’). Sensitivity within the direct spatial footprint of this pressure is therefore ‘High’. Although no specific evidence is described, confidence in this assessment is ‘High’ due to the incontrovertible nature of this pressure.  

None Very Low High
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

The species characterizing this biotope occur within the upper layers of the soft sediment (JNCC, 2015). This biotope does not occur on artificial substrata or rock habitats, although patches of sediment may support some of the species associated with this biotope. Any substratum other than the sediments on which this biotope is found would therefore lead to a loss of this biotope. Consequently, biotope resistance is assessed as 'None’, resilience is assessed as ‘Very Low’ (as the change, at the benchmark is permanent), and biotope sensitivity is assessed as ‘High’. 

None Very Low High
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

This biotope is found in medium to very fine gravelly sands  (Connor et al., 2004). The biotope would not be considered sensitive to a change to mixed sediments for muddy sands, as the sediment would still be likely to support the biotope without a fine sand fraction could, however, lead to a reduction in habitat suitability for both Lanice conchilega and the associated burrowing polychaetes.

An assessment of distribution records of Chaetozone setosa in the North Sea concluded that the species is usually associated with fine sediments (Chambers et al., 2007). Magelona spp. occur in coarse clean sand and fine clean sand (Rayment, 2007b). The polychaete Scoloplos armiger has relatively broad sediment preferences. Changes in sediment composition that alter the grade of sediment this species must move through can affect the suitability of the habitat. An increase in coarse composition to gravels would be expected to negatively impact this burrowing species. 

Sensitivity assessment. Changes in sediment that lead to increased finer sediments are likely to be tolerated, however, a change in the sediment Folk class classification at the pressure benchmark to gravel is likely to alter the abundance of Lanice conchilega and the composition of the species assemblage, resulting in loss of this biotope. Resistance is therefore assessed as 'None' and resilience as 'Very Low' (as the change at the pressure benchmark is permanent). The biotope is considered to have 'High' sensitivity to this pressure.

None Medium Medium
Q: High
A: High
C: High
Q: High
A: Low
C: High
Q: High
A: Low
C: High

Sedimentary communities are likely to be highly intolerant of substratum removal, which will lead to partial or complete defaunation, exposure of underlying sediment which may be anoxic and/or of a different character or bedrock and lead to changes in the topography of the area (Dernie et al., 2003). Any remaining species, given their new position at the sediment/water interface, may be exposed to conditions to which they are not suited, i.e. unfavourable conditions. Newell et al. (1998) state that removal of 0.5 m depth of sediment is likely to eliminate benthos from the affected area.

Removal of 30 cm of sediment will remove species that occur at the surface and within the upper layers of sediment. The extraction of sediment would remove the characterizing species Lanice conchilega, and the associated species present, including Cerastoderma edule, which is found to a depth of 5 cm and Nephtyid species and other polychaetes, such as Scoloplos armiger and Pygospio elegans that burrow between 5 and 20 cm into the sediment (Schӧttler, 1982; Pedersen, 1991; Kruse et al., 2004).   

Hydrodynamics and sedimentology (mobility and supply) influence the recovery of soft sediment habitats (Van Hoey et al., 2008). Recovery of the sedimentary habitat would occur via infilling, some recovery of the biological assemblage may take place before the original topography is restored, if the exposed, underlying sediments are similar to those that were removed.

Sensitivity assessment. Extraction of 30 cm of sediment will remove the characterizing biological component of the biotope. Resistance is assessed as ‘None’ and biotope resilience is assessed as ‘Medium’ as some sediment recovery may be required before recovery processes begin. Biotope sensitivity is therefore assessed as ‘Medium’. 

High High Not sensitive
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

The key characterizing species of this biotope (Lanice conchilega) has robust, flexible tubes and may retract below the surface. They are also able to rapidly rebuild or repair tubes (Nickolaidou, 2003). These characteristics reduce exposure to this pressure and enhance recovery. Rabaut et al. (2008) studied fisheries impacts at the species level in temperate sandy bottom areas. A controlled field manipulation experiment was designed focusing on areas with high densities of Lanice conchilega (i.e. Lanice conchilega reefs). A treatment zone was exposed to a one-off experimental trawling and the impact on, and recovery of the associated fauna was investigated for a period of 9 days post-impact. Community analysis showed a clear impact on associated species such as Eumida sanguinea, followed by a relatively quick recovery. The passage of a single beam trawl did not significantly alter the density of Lanice conchilega (Rabaut et al., 2008).

The characterizing polychaetes Magelona mirabilis, Chaetozone setosa and Spiophanes bombyx are soft bodied organism which will be exposed to abrasion while feeding at the sediment surface while feeding. Abrasion would be likely to cause physical damage to these species.

Juveniles and adults of Scoloplos armiger stay permanently below the sediment surface, and freely move without establishing burrows. While juveniles are only found a few millimetres below the sediment surface, adults may retreat to 10 cm depth or more (Reise, 1979; Kruse et al., 2004). The egg cocoons are laid on the surface and hatching time is 2-3 weeks during which these are vulnerable to surface abrasion.

Sensitivity assessment. The experiments by Rabaut et al. (2008) suggest that Lanice conchilega has ‘High’ resistance to abrasion, however, other associated species may be more impacted but species may be able to repair and recover from damage. Biotope resistance to a single abrasion event is assessed as ‘High’ based on the key characterizing species Lanice conchilega. There may be some damage to Lanice conchilega tubes and some reduction in abundances of the associated polychaete species but this unlikely to significantly alter the character of the biotope. Recovery from impacts of associated species is predicted to be ‘High’ and the biotope is assessed as ‘Not sensitive’.  

High High Not sensitive
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

The characterizing species of this biotope (Lanice conchilega) has robust, flexible tubes and may retract below the surface. They are also able to rapidly rebuild or repair tubes (Nickolaidou, 2003). These characteristics reduce exposure to this pressure and enhance recovery.

Rabaut et al. (2008) studied fisheries impacts at the species level in temperate sandy bottom areas. A controlled field manipulation experiment was designed focusing on areas with high densities of Lanice conchilega (i.e. Lanice conchilega reefs). A treatment zone was exposed to a one-off experimental trawling, and the impact on and recovery of the associated fauna was investigated for a period of 9 days post-impact. Community analysis showed a clear impact followed by a relatively quick recovery. The passage of a single beam trawl did not significantly alter the density of Lanice conchilega. Rabaut et al. (2009) also studied the direct mortality of Lanice conchilega as a consequence of sustained physical disturbance at varying frequencies to reflect the effect of beam trawl fisheries. Research was based on a laboratory experiment in which four different disturbance regimes were applied (disturbance every other 12, 24 and 48 h and no fishing disturbance as a control). Survival dropped significantly after 10 and 18 days (with a disturbance frequency of every 12 and 24 h, respectively). The results indicate that Lanice conchilega is relatively resistant to physical disturbance but that reef systems can potentially collapse under continuous high frequency disturbance.

Sparks-McConkey & Watling (2001) identified Chaetozone setosa as a common species that declined in abundance in response to experimental trawling. Tuck et al. (1998) found that following trawl disturbance, abundances of Chaetozone setosa had recovered and became greater at treatment sites than undisturbed sites 10 months after disturbance. Scoloplos armiger, however, had declined at disturbed sites.

Direct mortality (percentage of initial density) of Scoloplos armiger from a single pass of a beam trawl was estimated from experimental studies on sandy and silty grounds as 22% and 28% respectively. Mortality of Magelona sp. was estimated as 30% (Bergman & Van Santbrink, 2000a). 

Experimental intertidal dredging for cockles reduced the abundance of Scoloplos armiger in disturbed plots compared to control sites. These differences persisted for 56 days (Hall & Harding, 1997). Ferns et al. (2000) reported a decline of 31% in intertidal populations of Scoloplos armiger  in muddy sands when a mechanical tractor towed harvester was used (in a cockle fishery) (surpassing the study monitoring timeline). Scoloplos armiger demonstrated recovery >50 days after harvesting in muddy sands. 

Sensitivity assessment. The experiments by Rabaut et al. (2008, 2009) suggest that Lanice conchilega has ‘High’ resistance to single abrasion and penetration events. Other polychaetes within the biotope may be more impacted. Biotope resistance to a single event is assessed as ‘Medium’ based on the key characterizing species Lanice conchilega and associated characterizing species.  There may be some damage to tubes but this is expected to be sub-lethal and tubes are likely to be rapidly repaired. Recovery from impacts of associated species is predicted to be ‘High’ and the biotope is assessed as ‘Not sensitive’.  

High High Not sensitive
Q: Low
A: NR
C: NR
Q: High
A: High
C: High
Q: Low
A: Low
C: 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. 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. A decrease in suspended organic particles may reduce food supply to Lanice conchilega, however, this species can switch from suspension to deposit feeding and vice versa (Buhr & Winter, 1977). Supply of materials needed to build the tubes may be affected with a decrease in suspended solids but wave action would be likely to continue to re-suspend and transport sediments.

Sensitivity assessment. A decrease in turbidity and hence increased light penetration may result in increased phytoplankton production and hence increased food availability for suspension feeders, including Lanice conchilega and Cerastoderma edule. Therefore, reduced turbidity may be beneficial. In areas of high suspended sediment, a decrease may result in improved condition and recruitment due to a reduction in the clogging of filtration apparatus of suspension feeders and an increase in the relative proportion of organic particulates. However, a decrease in suspended organic particles in some areas may reduce food availability for deposit feeders resulting in lower growth or reduced energy for reproduction. 

Where increased turbidity results from organic particles then subsequent deposition may enhance food supply for suspension and deposit feeders within the biotope. Alternatively, if turbidity results from an increase in suspended inorganic particles then energetic costs may be imposed on these species as sorting and feeding becomes less efficient, reducing growth rates and reproductive success. Lethal effects are considered unlikely given the occurrence of Lanice conchilega and Cerastoderma edule and other associated species in estuaries where turbidity is frequently high from suspended organic and inorganic matter. Resistance and resilience are therefore assessed as 'High' and the biotope is considered to be 'Not sensitive'.

High High Not sensitive
Q: High
A: High
C: High
Q: High
A: High
C: High
Q: High
A: High
C: High

The tube of Lanice conchilega protrudes above the sediment by 1–4 cm, an adaptation to trap suspended particles (Zühlke, 2001). Furthermore, Ziegelmeier (1952) showed that the tubes of Lanice conchilega increased the height with increasing sedimentation so that it could continue feeding and metabolizing aerobically. This enables Lanice conchilega to switch between deposit and suspension feeding (Buhr, 1976). Therefore, Lanice conchilega is suitably adapted to discrete events of sedimentation, characteristic of sandy and muddy sand intertidal flats wher epopulations of this species occur. Lanice conchilega  has dominated areas of Manila clam cultivation where protective netting had led to an increase in sedimentation (Spencer et al., 1998) and in areas where oysters were cultivated and sedimentation increased (Sylvand, 1995).

Chaetezone setosa occurs in areas subject to high natural rates of sedimentation where other benthic macrofauna were excluded (Wlodarska-Kowalczuk et al., 2007) although this is likely to be due to rapid recolonization rather than survival. In general, the surface deposit feeders, Chaetozone setosa, Magelona mirabilis and Spiophanes bombyx will be smothered by a depoist at the benchmark and prevented from feeding while this remains in place. No evidence was found on the ability of these species to reposition within fine sediment deposits or survive burial.

Bijkerk (1988, results cited from Essink, 1999) indicated that the maximal overburden through which Scoloplos could migrate was 50 cm in sand and mud. No further information was available on the rates of survivorship or the time taken to reach the surface. Warner (1971) simulated the effects of dredge disposal of different thicknesses on animals in aquaria or plastic cores for 2 weeks. In core experiments at temperatures ranging from 14 to 18°C and 20 to 21°C, there was a relationship between vertical migration distance and sediment depth for the congener Scoloplos fragilis. This species could vertically migrate through 30 cm of sand. In other core experiments in silt-clay at temperatures of 17°C to 18°C, there was a suggestion of reduced efficiency of burrowing in finer grained sediment where even the smallest amount of silt-clay proportion tested (20%) affected the burrowing ability of this species.

Sensitivity assessment. Based upon the ability of Lanice conchilega to increase tube elevation above the sediment surface when deposition has increased, it is likely that Lanice conchilega is resistant to the effects of a single discrete event of deposition of 5 cm. Although Scoloplos armiger is considered to be able to migrate vertically, this may be limited where the overburden consists of fine sediments (based on Maurer et al., 1978). Sensitivity to continuous events will depend on tidal hydrodynamics, and spawning and recruitment, prior to the event (see recover/resilience rates). As the benchmark refers to a single discrete event, biotope resistance is assessed as ‘High’ based on Lanice conchilega and resilience is ‘High’, therefore this biotope is ‘Not sensitive’.

Low High Low
Q: Low
A: NR
C: NR
Q: High
A: Low
C: High
Q: Low
A: Low
C: Low

No evidence was found to assess sensitivity of Lanice conchilega to heavy sedimentation. It is likely that intertidal flats may sometimes be subject to sedimentation following storm events but no species-specific examples were found. Ziegelmeier (1952) showed that Lanice conchilega increased the height of its tube top with increasing sedimentation so that it could continue feeding and respire. However, heavy sedimentation of 30 cm during a single event may have a severe effect on Lanice conchilega and the associated species.

Bijkerk (1988, results cited from Essink, 1999) indicated that the maximal overburden through which Scoloplos could migrate was 50 cm in sand and mud. No further information was available on the rates of survivorship or the time taken to reach the surface. Warner (1971) simulated the effects of dredge disposal of different thicknesses on animals in aquaria or plastic cores for 2 weeks. In core experiments at temperatures ranging from 14 to 18°C and 20 to 21°C, there was a relationship between vertical migration distance and sediment depth for the congener Scoloplos fragilis. This species could vertically migrate through 30 cm of sand. In other core experiments in silt-clay at temperatures of 17°C to 18°C, there was a suggestion of reduced efficiency of burrowing in finer grained sediment where even the smallest amount of silt-clay proportion tested (20%) affected the burrowing ability of this species.

 Sensitivity assessment. Although Scoloplos armiger is considered to be able to migrate vertically, this may be limited where the overburden consists of fine sediments (based on Maurer et al., 1978). The addition of 30 cm of sediment would prevent the surface deposit feeders from extending palps to the surface. It is unlikely that these species could emerge from a 30 cm thick deposit of fine materials., although some individuals may survive and sediment may be rapidly removed by tide and wave action. It is likely, however, that there would be considerable mortality of characterizing species, including Lanice conchilega. Resistance is assessed as ‘Low’ and resilience as ‘High’. Sensitivity is therefore assessed as ‘Low’. 

Not Assessed (NA) Not Assessed (NA) Not assessed (NA)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Not assessed.

No evidence (NEv) No Evidence (NEv) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No evidence.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No information was found concerning the intolerance of the biotope or the characterizing species to noise. The palps of polychaetes are likely to detect vibrations and would probably be withdrawn as a predator avoidance mechanism. However, it is unlikely that the biotope and characterizing species will be affected by noise or vibrations caused by noise at the level of the benchmark and this pressure is assessed as ‘Not relevant’.

No evidence (NEv) No Evidence (NEv) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No evidence. As this feature is not characterized by the presence of primary producers, it is not considered that shading would alter the character of the habitat directly.

High High Not sensitive
Q: Low
A: NR
C: NR
Q: High
A: High
C: High
Q: Low
A: Low
C: Low

Barriers that reduce the degree of tidal excursion may alter larval supply to suitable habitats from source populations. Conversely, the presence of barriers at brackish waters may enhance local population supply by preventing the loss of larvae from enclosed habitats to environments, which are unfavourable, reducing settlement outside of the population. Resistance to this pressure is assessed as 'High' and resilience as 'High' by default. This biotope is therefore considered to be 'Not sensitive'.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

‘Not relevant’ to seabed habitats. NB. Collision by grounding vessels is addressed under ‘surface abrasion’.

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

'Not relevant'.

Biological Pressures

 ResistanceResilienceSensitivity
Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Characterizing and associated species are not cultivated or transplanted and this pressures is considered 'Not relevant'.

No evidence (NEv) No Evidence (NEv) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No evidence.

No evidence (NEv) No Evidence (NEv) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No evidence was found for microbial pathogens, parasites or diseases of Lanice conchilega or other characterizing polychaetes. 

Not relevant (NR) Not relevant (NR) Not relevant (NR)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

The direct, physical impacts of removal are assessed through the abrasion and penetration of the seabed pressures. The sensitivity assessment for this pressure considers any biological/ecological effects resulting from the removal of target species on this biotope. No characterizing or associated benthic species are targeted by recreational or commercial fishers and this pressure is considered to be 'Not relevant'.

 

Low High Low
Q: High
A: High
C: High
Q: High
A: Medium
C: High
Q: High
A: Medium
C: High

Species within this biotope may be removed or damaged by static or mobile gears that are targeting other species. These direct, physical impacts are assessed through the abrasion and penetration of the seabed pressures. The sensitivity assessment for this pressure considers any biological/ecological effects resulting from the removal of non-target species on this biotope.

Lanice conchilega (sand mason worm) is the dominant polychaete in this biotope. It qualifies as an ‘ecosystem engineer’ in that it changes and/or creates a habitat, which affects the abundance of other species (Jones et al., 1994, 1997). The tube which Lanice conchilega builds provides structure and stabilizes the sediment (Jones & Jago, 1993). The tubes also obstruct the activities of predatory burrowers enabling other sedentary animals to establish themselves (Wood, 1987). The burrows and tubes allow oxygenated water to penetrate into the sediment, the oxic upper layer of sediment to support shallow burrowing species such as amphipods (Ampelisca spp., Bathyporeia spp. & Gammarus spp.) and small Crustacea. Lanice conchilega may therefore increase species richness and abundance through habitat modification (Rabaut et al., 2007; Zuhlke, 2001). Dense aggregates of Lanice conchilega have the potential to enhance food supply to birds and fish due to increased biodiversity (Rabaut et al., 2010; Godet et al., 2008; Van Hoey et al., 2008; Callaway et al., 2010; De Smet et al., 2013).

Sensitivity assessment. The removal of non-target species may result in changes to the biological community and hence the classification of the assemblage type as assessed in the biotope. Incidental removal of Lanice conchilega would alter the biotope classification and most likely result in changes to the habitat and associated species. As Lanice conchilega is sedentary and present at the surface, resistance is ‘Low’ and resilience is ‘High’. Biotope sensitivity is therefore assessed as ‘Low’. This assessment considers the ecological effect of removal, the abrasion pressure assessment provides evidence for the robustness of tubes and the species may be resistant to removal by bottom trawls deployed at low intensity (e.g. single passes).

Importance review

Policy/Legislation

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

Exploitation

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

Additional information

-

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Citation

This review can be cited as:

McQuillan, R. M. & Tillin, H,M. 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: http://www.marlin.ac.uk/habitat/detail/116

Last Updated: 28/06/2006