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information on the biology of species and the ecology of habitats found around the coasts and seas of the British Isles

Ampelisca spp., Photis longicaudata and other tube-building amphipods and polychaetes in infralittoral sandy mud

20-03-2020

Summary

UK and Ireland classification

UK and Ireland classification

Description

Sublittoral stable cohesive sandy muds occurring over a wide depth range may support large populations of semi-permanent tube-building amphipods and polychaetes. In particular large numbers of the amphipods Ampelisca spp. and Photis longicaudata may be present along with polychaetes such as Lagis koreni. Other important taxa may include bivalves such as Nucula nitidosaChamelea gallina, Abra alba and Kurtiella bidentata and the echinoderms Echinocardium cordatum and Amphiura brachiata. In some areas, polychaetes such as Spiophanes bombyx and Polydora ciliata may also be conspicuously numerous. This community is poorly known, appearing to occur in restricted patches. In some areas, it is possible that AmpPlon may develop as a result of moderate organic enrichment. A similar community in mud has also been reported in the Baltic that is characterized by large populations of amphipods such as Ampelisca spp., Corophium spp. and Haploops tubicola (see Petersen, 1918; Thorson, 1957) and it is not known if SMU.AmpPlon is a UK variant of this biotope.

In some areas of the Irish Sea, this biotope is reported to be a temporal variant of AalbNuc, SsubNhom, and LkorPpel. Some researchers consider these biotopes to be part of a wider muddy sand community that varies temporally depending on changes in sediment deposition and recruitment as was reported in areas of Red Wharf Bay off the Welsh coast (E.I.S. Rees pers. comm. 2002). (Information from Connor et al., 2004; JNCC, 2015). 

Depth range

0-5 m, 5-10 m, 10-20 m, 20-30 m

Additional information

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Sensitivity reviewHow is sensitivity assessed?

Sensitivity characteristics of the habitat and relevant characteristic species

SS.SMu.ISaMu.AmpPlon is a sublittoral biotope characterized by a tube mat created by large numbers of the tube-building amphipods, Ampelisca spp. and Photis longicaudata although tube building polychaetes such as Polydora ciliata and Spiophanes bombyx also occur (Connor et al., 2004). It may be similar to the amphipod dominated 'Boreal-Arctic Ampelisca' community described in the Baltic by Thorson (1957) and the Ampelisca dominated community in Massachusetts, USA, described by Mills (1967, 1969).  Mills (1967, 1969) reported that high numbers of Ampelisca colonized sand falts on Barnstable Harbour, Massachusetts in 1960-62.  The resultant tube mat was patchy but often covered hundreds of square metres, and the density of individuals was great even in winter.  The tube mat also increased the surface complexity of the sandflat, provided surface area for benthic diatoms, and increased the abundance of infaunal polychaetes. In Maine, the amphipod tube mat resulted in an increase in diversity in the previously species-poor habitat.  Filter-feeding by Ampelisca and the resultant faeces trapped organic particulates at the surface, which in turn increased the organic content between the amphipod tubes which favoured the selective deposit-feeding polychaetes. However, it also decreased the median sedimentary particle size, which caused instability so that minor breaks in the mat were sufficient to cause wash out of the tube mat, especially on windy days, or storms (Mills, 1967, 1969). Mills (1967, 1969) reported that egg-bearing females settled in uncolonized areas so that juveniles avoided competition from remaining adults in the tube mat and colonized areas that were more physically stable. As a result, they abandoned old areas and colonized new spaces. Mills (1969) concluded that the Ampelisca community was adapted to a 'dynamic instability' and frequent changes in the area it occupied.  However, Mackenzie et al. (2006) reported that dense Ampelisca tube mat persisted for at least five years in the deeper waters sheltered from wave action in Raritan Bay, New Jersey. Therefore, it is probably a transient and patchy community, which may be a transient overlay over other biotopes, such as the AalbNuc, SsubNhom and LkorPpel biotopes in the Irish Sea (see Connor et al., 2004). 

Therefore, the tube-building amphipods such as Ampelisca spp. and Photis longicaudata are probably the key structural species in the biotope and are the main focus of the sensitivity assessment, as loss of the tube mat would result in the loss of the biotope. The sensitivity of the other tube-builders such as Polydora ciliata and Spiophanes bombyx are discussed where relevant. 

Resilience and recovery rates of habitat

The amphipod genus Ampelisca has life history traits that allow them to recovery quickly where populations are disturbed. They do not produce large numbers of offspring but reproduce regularly and the larvae are brooded, giving them a higher chance of survival within a suitable habitat than free-living larvae. Ampelisca has a short lifespan and reaches sexual maturity in a matter of months allowing a population to recover abundance and biomass in a very short period (MES, 2008). Experimental studies have shown Ampelisca abdita to be an early colonizer, in large abundances of defaunated sediments where local populations exist to support recovery (McCall, 1977) and Ampelisca abdita have been shown to migrate to, or from, areas to avoid unfavourable conditions (Mills, 1967, 1969; Nichols & Thompson, 1985). Ampelisca spp. are very intolerant of oil contamination and the recovery of then Ampelisca populations in the fine sand community in the Bay of Morlaix took up to 15 years following the Amoco Cadiz oil spill, probably due to the amphipods' low fecundity, lack of pelagic larvae and the absence of local unperturbed source populations (Poggiale & Dauvin, 2001). Mills (1967) reported that Ampelisca vadorum and Ampelisca abdita produced only one brood per generation but that there were two or more generations per year. In the English Channel, two reproductive patterns were identified. Species such as Ampelisca tenuicornis and Ampelisca typica produced two generations per year. The juveniles born in May-June were able to brood in September-October (Dauvin, 1988b; Dauvin, 1988c). Species such as Ampelisca armoricana and Ampelisca sarsi produced only one brood per generation and per year (Dauvin, 1989; Dauvin, 1988d). Ampelisca brevicornis showed an intermediate cycle with one generation per year during cold years (cold spring) and two generations per year during warm years (warm spring) and its cycle is intermediate between univoltine cycle and bivoltine cycle (Dauvin, 1988b,c,d,e; Dauvin, 1989; Dauvin & Bellan-Santini, 1990).  In addition, Mills (1967, 1969) concluded that Ampelisca was a 'vagrant species', adapted to frequent changes in the area occupied and the ability to rapidly colonize new habitat due to the tendency of egg-bearing or gravid females to relocate via the plankton, and its short generation time resulting in dramatic increases in numbers in colonized areas. However, Mackenzie et al. (2006) reported that dense Ampelisca tube mat persisted for at least five years in deeper waters sheltered from wave action in Raritan Bay, New Jersey.

Polydora is a small, sedentary, burrowing polychaete worm up to 3 cm long. All Polydora spp. make a U-shaped tube from small particles (Hayward & Ryland, 1995b). Polydora ciliata usually burrows into substrata containing calcium carbonate such as limestone, chalk and clay, as well as shells or oysters, mussels and periwinkles (Fish & Fish, 1996). The sexes are separate and breeding has been recorded in spring in a number of locations. In northern England, it has been recorded to occur from February until June and three or four generations succeed one another during the spawning period (Gudmundsson, 1985). Eggs are laid in a string of capsules that are attached by two threads to the wall of the burrow (Fish & Fish, 1996). After a week the larvae emerge and are believed to have a pelagic life of 2-6 weeks before settling. Length of life is no more than one year (Fish & Fish, 1996). Almeda et al. (2009) suggested low filtration rates and low growth rates despite high food availability for Polydora ciliata larvae, which suggested a compromise to ensure efficient larval dispersion. Larvae are substratum specific, selecting rocks according to their physical properties or selecting sediment depending on particle size. Larvae of Polydora ciliata have been collected as far as 118 km offshore (Murina, 1997). Adults of Polydora ciliata produce a 'mud' resulting from the perforation of soft rock substrata and the larvae of the species settle preferentially on substrata covered with mud (Lagadeuc, 1991). The settling of the first generation in April is followed by the accumulation and active fixing of mud continuously up to a peak during May. The following generations do not produce a heavy settlement due to interspecific competition and heavy mortality of the larvae (Daro & Polk, 1973). The tubes built by Polydora sometimes agglomerate to form layers of mud up to an average of 20 cm thick. Later in the year, the surface layer cannot hold the lower layers of the mud mat in place. They crumble away and are then swept away by water currents. The empty tubes of Polydora may saturate the sea in June. The early reproductive period of Polydora ciliata often enables the species to be the first to colonize available substrata (Green, 1983). For example, in colonization experiments in Helgoland (Harms & Anger, 1983), Polydora ciliata settled on panels within one month in the spring.

Other polychaetes in the biotope are likely to also recolonize disturbed areas rapidly. For example, Spiophanes (e.g. Spiophanes bombyx) have opportunistic life strategies and exhibit small size, rapid maturation and a short lifespan of 1-2 years with the production of large numbers of small propagules. Two years after dredging, abundances of opportunistic species were generally elevated relative to pre-dredging levels while communities had become numerically dominated (50-70 %) by Spiophanes bombyx (Gilkinson et al., 2005). Van Dalfsen et al. (2000) found that polychaetes recolonized a dredged area within 5-10 months (reference from Boyd et al., 2005), with biomass recovery predicted within 2-4 years. Spiophanes bombyx is regarded as a typical 'r' selecting species with a short lifespan, high dispersal potential and high reproductive rate (Niermann et al., 1990). It is often found at the early successional stages of variable, unstable habitats that it is quick to colonize following a perturbation (Pearson & Rosenberg, 1978). Its larval dispersal phase may allow the species to colonize remote habitats. McLusky et al. (1983) examined the effects of bait digging on blow lug populations in the Forth Estuary. Dug and infilled areas and unfilled basins left after digging repopulated within one month, whereas mounds of dug sediment took longer and showed a reduced population. Basins accumulated fine sediment and organic matter and showed increased population levels for about 2-3 months after digging. Overall recovery is generally regarded as rapid. Pygospio elegans were significantly depleted for >100 days after harvesting (surpassing the study monitoring timeline) and Scoloplos armiger demonstrated recovery >50 days after harvesting in muddy sands (Ferns et al., 2000). In summary, these studies suggest recovery from fisheries pressures occurs in four months to >3 years depending upon the harvesting method (such as hand digging or mechanical dredging) and the size of the area impacted. As a tube building polychaete, Pygospio elegans aids stabilisation of sediments following disturbance. Recolonization and hence recovery may be aided by bedload transport of juvenile polychaetes and bivalves. Recolonization of Pygospio elegans was observed in two weeks by Dittmann et al. (1999) following one-month long defaunation of the sediment.

Resilience assessment. Removal of the characterizing species and the 'tube mat' would result in the biotope being lost and/or reclassified. Amphipods brood their young so that dispersal is limited to local movements of these sub-juveniles and migration of the adults and hence recruitment is limited by the presence of local, unperturbed source populations (Poggiale & Dauvin, 2001). Dispersal of sub-juveniles may be enhanced by the brooding females leaving their tubes and swimming to uncolonized areas of substratum before the eggs hatch (Mills, 1967, 1969). Nevertheless, Ampelicsca is an early colonizer and brooding of juveniles allows for rapid localized population growth. The tube building polychaetes such as Polydora ciliata generally disperse via pelagic larvae (Fish & Fish, 1996) and, therefore, recruitment may occur from distant populations. These are likely to recolonize disturbed areas first, although the actual pattern will depend on the recovery of the habitat, season of occurrence and other factors.  Therefore, where perturbation removes a portion of the population or even causes local extinction (resistance High, Medium or Low) resilience is likely to be 'High' (<2 years) as long as recruitment from neighbouring areas and/or adult migration is possible. However, even in areas of suitable habitat that are isolated, where total extinction of the population occurs (resistance 'None') recovery is likely to depend on favourable hydrodynamic conditions that will allow recruitment from farther away. However, once an area has been recolonized, restoration of the biomass of the characterizing species is likely to occur quickly and resilience is likely to be 'Medium' (full recovery within 2-10 years).

NB: The resilience and the ability to recover from human induced pressures is a combination of the environmental conditions of the site, the frequency (repeated disturbances 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 before 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

Use / to open/close text displayedResistanceResilienceSensitivity
High High Not sensitive
Q: Medium
A: Low
C: Medium
Q: High
A: High
C: High
Q: Medium
A: Low
C: Medium

Amphipods were reported to have a low tolerance to temperature changes (Bousfield, 1973) although lethal limits were not given. However, the amphipods that occur within this habitat are mobile and can avoid unfavourable conditions to some extent. Ampelisca brevicornis is widespread in the North East Atlantic but is also recorded from the White Sea, south to the Iberian Peninsula and the Mediterranean, and as far as the coasts of South Africa, India, China and the Korean Peninsula (OBIS, 2020). It has been recorded in areas where sea surface temperatures (SST) range from 5 to 30°C, although most records occur in the 10-15°C range (OBIS, 2020).  Similarly, Ampelisca typica is also recorded from the White Sea, through the North East Atlantic to the Iberian Peninsula, and in areas where the SST ranged from 5-25°C although most records occurred in 10-15°C (OBIS, 2020). However, other species, e.g. Ampelisca sarsi have more limited distribution south of the British Isles (OBIS, 2020). Photis longicaudata is also widespread in the North East Atlantic but is also recorded from the Barents Sea, south to the Iberian Peninsula and the Mediterranean, and as far as the coasts of South Africa, India, China and the Korean Peninsula, and in the west Atlantic from Florida south to Uruguay and in areas where the SST ranged from minus 5-0°C and 5-30°C although most records occurred in 10-15°C (OBIS, 2020).

Murina (1997) categorized Polydora ciliata as a eurythermal species because of its ability to spawn in temperatures ranging from 10.6-19.9°C. This is consistent with a wide distribution in north-west Europe, which extends into the warmer waters of Portugal and Italy (Pardal et al., 1993; Sordino et al., 1989). In the western Baltic Sea, Gulliksen (1977) recorded high abundances of Polydora ciliata at temperatures of 7.5 to 11.5°C and in Whitstable, in Kent, where sea temperatures varied between 0.5 and 17°C (Dorsett, 1961). Growth rates may increase if the temperature rises. For example, at Whitstable in Kent, Dorsett (1961) found that a rapid increase in growth coincided with the rising temperature of the seawater during March. No information was found regarding the intolerance of Spiophanes bombyx to temperature. Spiophanes bombyx is found in the Mediterranean (Hayward & Ryland, 1995b), which is likely to be warmer than the waters around Britain and Ireland.

Sensitivity assessment. Typical surface water temperatures around the UK coast vary seasonally from 4-19°C (Huthnance, 2010). The characteristic tube mat forming species are distributed to the north and south of the British Isles and are likely to be able to resist a long-term increase in temperature of 2°C and may resist a short-term increase of 5°C in UK waters. Resistance and resilience are, therefore, assessed as 'High' and the biotope assessed as 'Not Sensitive' at the benchmark level.

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

Amphipods were reported to have a low tolerance to temperature changes (Bousfield, 1973) although lethal limits were not given. However, the amphipods that occur within this habitat are mobile and can avoid unfavourable conditions to some extent. Ampelisca brevicornis is widespread in the North East Atlantic but is also recorded from the White Sea, south to the Iberian Peninsula and the Mediterranean, and as far as the coasts of South Africa, India, China and the Korean Peninsula (OBIS, 2020). It has been recorded in areas where sea surface temperatures (SST) range from 5 to 30°C, although most records occur in the 10-15°C range (OBIS, 2020).  Similarly, Ampelisca typica is also recorded from the White Sea, through the North East Atlantic to the Iberian Peninsula, and in areas where the SST ranged from minus 5-0°C and 5-25°C although most records occurred in 10-15°C (OBIS, 2020). However, other species, e.g. Ampelisca sarsi have more limited distribution south of the British Isles (OBIS, 2020). Photis longicaudata is also widespread in the North East Atlantic but is also recorded from the Barents Sea, south to the Iberian Peninsula and the Mediterranean, and as far as the coasts of South Africa, India, China and the Korean Peninsula, and in the west Atlantic from Florida south to Uruguay and in areas where the SST ranged from minus 5-0°C and 5-30°C although most records occurred in 10-15°C (OBIS, 2020).

Murina (1997) categorized Polydora ciliata as a eurythermal species because of its ability to spawn in temperatures ranging from 10.6-19.9°C. This is consistent with a wide distribution in north-west Europe. In the western Baltic Sea, Gulliksen (1977) recorded high abundances of Polydora ciliata at temperatures of 7.5 to 11.5°C and in Whitstable, Kent, abundance was high when winter water temperatures dropped to 0.5°C (Dorsett, 1961). During the extremely cold winter of 1962/63, Polydora ciliata was unaffected (Crisp, 1964).  Spiophanes bombyx is found in waters off Denmark (Thorson, 1946), which are likely to be colder than British and Irish waters.

Sensitivity assessment. Typical surface water temperatures around the UK coast vary seasonally from 4-19°C (Huthnance, 2010). The characteristic tube mat forming species are distributed to the north and south of the British Isles and are likely to be able to resist a long-term decrease in temperature of 2°C and may resist a short-term decrease of 5°C in UK waters. Resistance and resilience are, therefore, assessed as 'High' and the biotope assessed as 'Not Sensitive' at the benchmark level.

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

Monitoring at a Spanish desalination facility where discharges close to the outfall reached a salinity of 53, found that amphipods, including Ampelisca spp. were sensitive to the increased salinity and that species free-living in the sediment were most sensitive (De-la-Ossa-Carretero et al., 2016).  Roberts et al. (2010b) reported that the effects of brine discharge were dependent on the receiving environment but that the effects were limited to with 10s of metres of the outfall. In their review, they reported that polychaete abundance and diversity decreased adjacent to a brine outfall in Alicante, Spain and that the family Ampharaetidae were the most sensitive while the family Paraonidae were the least sensitive (Ruso et al., 2008 cited in Roberts et al., 2010b).  Polydora ciliata is a euryhaline species inhabiting fully marine and estuarine habitats. It is unlikely that Spiophanes bombyx would experience hypersaline conditions, therefore unlikely to be adapted to such conditions.

Sensitivity assessment. The characterizing species of this biotope are euryhaline and likely to be resistant to increases in salinity. However, the biotope occurs full saline conditions (Connor et al., 2004) and is unlikely to experience hypersaline conditions. Therefore, exposure to hypersaline effluent (>40) might result in the death of a portion of the individuals in the population, especially Ampelisca and some polychaetes species. Therefore, resistance is assessed as 'Low' but with 'Low' confidence. Resilience is likely to be High, so the biotope is assessed as 'Low' sensitivity to an increase in salinity at the pressure benchmark.

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

Ampelisca brevicornis was recorded from areas where the sea surface salinity (SSS) ranged between 5 and 35 psu although most records were in the range 15-35 psu and Ampelisca typica was recorded from areas where the SSS ranged between 15 and 35 psu although most records were in the range 30-35 psu (OBIS, 2020). Photis longicaudata was recorded from areas where the sea surface salinity ranged between 10 and 35 psu although most records were in the range 30-35 (OBIS, 2020).

Polydora ciliata is a euryhaline species inhabiting fully marine and estuarine habitats. In an area of the western Baltic Sea, where bottom salinity was between 11.1 and 15.0 psu Polydora ciliata was the second most abundant species with over 1000 individuals per m2 (Gulliksen, 1977). Spiophanes bombyx is a euryhaline species (Bailey-Brook, 1976; Maurer & Lethem, 1980), inhabiting fully saline and estuarine habitats.

Sensitivity assessment. SS.SMu.ISaMu.AmpPlon occurs in full salinity conditions (Connor et al., 2004). The evidence from distribution records (OBIS, 2020) suggests that the charactersitic amphipods vary in salinity tolerance but prefer full salinity conditions, except perhaps Ampelisca brevicornis, whereas the typical polychaetes are euryhaline. Therefore, a change in salinity regime from 'full' (30-35) to 'reduced' (18-30) might result in a reduction in the amphipod abundance and degradation of the tube mat. Hence, resitance is assessed as 'Medium' to represent the possible loss of a proportion of the resident community but with 'Low' confidence.  Resilience is probably 'High' so the biotope sensitivity is assessed as 'Low' to a decrease in salinity at the benchmark level.

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

Changes in water flow are likely to change the sediment characteristics in which the biotope occurs, primarily by resuspending and preventing deposition of finer particles (Hiscock, 1983). A decrease in water flow is unlikely to cause any impact on the biotope as species are adapted to incremental deposition, typical of low energy environments such as those where the sandy muds typical of this biotope occur. However, an increase would likely result in a decrease in tube-building material for the characterizing species, and the lack of deposition of particulate matter at the sediment surface would reduce food availability for the deposit feeders in the biotope. The resultant energetic cost over one year would be likely to result in some mortality of tube-builders and infauna. For example, Polydora ciliata was present and colonized test panels in Helgoland in three areas; two exposed to strong tidal currents and one site sheltered from currents (Harms & Anger, 1983). However, very strong water flows may sweep away Polydora colonies, often in a thick layer of mud on a hard substratum. 

The most damaging effect of increased flow rate would be the erosion of the medium to fine muddy sand substratum as this could eventually lead to loss of the habitat. Amphipods are thought to stabilize the intertidal sediments in which they reside (Mouritsen et al., 1998; MacKenzie et al., 2006). Mills (1967, 1969) noted that the tubes of Ampelisca increased the surface complexity of the sandflat in Barnstable Harbour, Massachusetts. However, their feeding decreased the median grain size of the sediment, towards fine particulates, and resulted in instability. Mills (1969) stated that minor breaks in the mat of Ampelisca tubes were sufficient to cause large areas to wash out, especially on windy days in a rising or falling tide, although no flow rates were given. Emergent species, such as the Polydora ciliata tubes that characterize this biotope, may create turbulent flow leading to particle resuspension. Additionally, where a change in water flow rate changes sediment characteristics, with increased deposits of coarser sediments, characterizing species may no longer be supported due to particular substratum preferences. 

Sensitivity assessment. SS.SMu.ISaMu.AmpPlon is recorded on sandy muds, which are typical of low energy environments i.e. wave sheltered and/or weak (<0.5 m/s) and very weak (negligible) tidal streams (Connor et al., 2004). Sand particles are likely to be eroded at about 0.20 m/s (based on Hjulström-Sundborg diagram, Sundborg, 1956), although the sandy muds are more cohesive and require higher flow rates to resuspend. Also, the tube mat may help to stabilize the sediment surface, initially, until itself destabilizes due to the build-up of finer particulates (Mills, 1967, 1969). Although a decrease in water flow rate is likely to be irrelevant, an increase in water flow at the pressure benchmark is may result in loss of parts of the characterizing mat of tubes formed by Ampelisca spp., and Polydora.  Furthermore, stronger currents (e.g. due to storms) are likely to wash away the community of semi-permanent tube-building amphipods and polychaetes that characterize the biotope. Therefore, resistance is assessed as 'Low' and resilience as 'High' so that sensitivity is assessed as 'Low' sensitivity to a change in water flow at the pressure benchmark level.

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

SS.SMu.ISaMU.AmpPlon is recorded from 0-30 m (Connor et al., 2004) and is predominately subtidal. Therefore, this pressure is probably 'Not relevant'.  

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

Where the biotope occurs in the shallow subtidal, it is likely to be affected by winter storms. Storms may cause dramatic changes in the distribution of macro-infauna by washing out dominant species, opening the sediment to recolonization by adults and/or available spat/larvae (Eagle, 1975; Rees et al., 1976; Hall, 1994) and by reducing the success of recruitment by newly settled spat or larvae (see Hall, 1994 for review). 

Feeding of the characterizing species may be impaired in strong wave action and changes in wave exposure may also influence the supply of particulate matter for tube-building polychaetes and amphipods. Mills (1967, 1969) noted that the tubes of Ampelisca increased the surface complexity of the sandflat in Barnstable Harbour, Massachusetts. Also, their feeding decreased the median grain size of the sediment, towards fine particulates, and resulted in instability. Mills (1969) stated that minor breaks in the mat of Ampelisca tubes were sufficient to cause large areas to wash out, especially on windy days in a rising or falling tide and reported that Ampelisca flats in Barnstable, Massachusetts, were damaged noticeably by winter storms (Mills, 1967, 1969). Mackenzie et al. (2006) reported that dense mats of Ampelisca persisted in Raritan Bay, New Jersey for at least five years because the mats occupied deeper waters (ca 7 m) and were sheltered from wave action. 

Sensitivity assessment. SS.SMu.ISaMu.AmpPlon is recorded on sandy muds, which are typical of low energy environments i.e. wave sheltered and/or weak (<0.5 m/s) and very weak (negligible) tidal streams (Connor et al., 2004). The tube mat that characterizes this biotope is probably very susceptible to damage or removal by during winter storms or on windy days, especially in the shallow examples of the habitat. Mills (1969) considered that the Ampelisca dominated tube mat demonstrated 'dynamic instability' due to its need to move from areas of physical disturbance and sediment modification.  Although the biotope occurs in low energy conditions (Connor et al., 2004) it is uncertain if a 3-5% change in significant wave height (the benchmark level) would be significant. Therefore, resistance is assessed as 'High', resilience as 'High' and sensitivity is assessed as 'Not Sensitive' at the benchmark level. 

Chemical Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
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

This pressure is Not assessed but any evidence is presented where available.

Experimental studies with various species suggest that polychaete worms are quite resistant to heavy metals (Bryan, 1984). Polydora ciliata occurred in an area of the southern North Sea polluted by heavy metals but was absent from sediments with very high heavy metal levels (Diaz-Castaneda et al., 1989). For most metals, toxicity to crustaceans increases with decreased salinity and elevated temperature, therefore marine species living within their normal salinity range may be less susceptible to heavy metal pollution than those living in salinities near the lower limit of their salinity tolerance (McLusky et al., 1986). In laboratory investigations, Hong & Reish (1987) observed 96-hour LC50 water column concentrations of between 0.19 and 1.83 mg/l for several species of amphipod. Corophium volutator is highly intolerant of metal pollution at levels often found in estuaries from industrial outfalls and contaminated sewage. A concentration of 38 mg Cu/l was needed to kill 50% of Corophium volutator in 96-hour exposures (Bat et al., 1998). Other metals are far more toxic to Corophium volutator, e.g. zinc is toxic over 1 mg/l and toxicity to metals increases with increasing temperature and salinity (Bryant et al., 1985). Mortality of 50% is caused by 14 mg/l (Bat et al., 1998). Although exposure to zinc may not be lethal, it may affect the perpetuation of a population by reducing growth and reproductive fitness. Mercury was found to be very toxic to Corophium volutator, e.g. concentrations as low as 0.1 mg/l caused 50% mortality in 12 days. Other metals known to be toxic include cadmium, which causes 50% mortality at 12 mg/l (Bat et al., 1998); and arsenic, nickel and chromium which are all toxic over 2 mg/l (Bryant et al., 1984; Bryant et al., 1985a; 1985).

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

This pressure is Not assessed but any evidence is presented where available.

In general, soft-sediment inhabitants, especially infaunal polychaetes, are particularly affected by oil pollution (Suchanek, 1993). For example, Jacobs (1980) investigated the effects of the Amoco Cadiz oil spill in 1978 and noted that the numbers of spionidae polychaetes decreased after the spill. In an analysis of kelp holdfast fauna following the Sea Empress oil spill in Milford Haven the fauna present, including Polydora ciliata, showed a strong negative correlation between numbers of species and distance from the spill (SEEEC, 1998). After the extensive oil spill in West Falmouth, Massachusetts, Grassle & Grassle (1974) followed the settlement of polychaetes in the disturbed area. Species with the most opportunistic life histories, including Polydora ligni, were able to settle in the area. This species has some brood protection which enables larvae to settle almost immediately in the nearby area (Reish, 1979). Furthermore, Gray et al. (1990) found that Scoloplos armiger was a dominant species in uncontaminated soft sediments at a case study site adjacent to the Ekofisk oil field but was not present at contaminated sites, suggesting Scoloplos armiger are also intolerant to hydrocarbon contaminates.

Amphipods in general and ampeliscid amphipods, in particular, seem particularly intolerant of contamination with oil. Dauvin (1998) reported reductions in abundance, biomass and production of Ampelisca sp. following the Amoco Cadiz oil spill. Furthermore, light fractions (C10 - C19) of oils are much more toxic to Corophium volutator than heavier fractions (C19 - C40). In exposures of up to 14 days, light fraction concentrations of 0.1 g/kg sediment caused high mortality. It took 9 g/kg sediment to achieve similar mortalities with the heavy fraction (Brils et al., 2002). In the Forth Estuary, Corophium volutator was excluded for several hundred metres around the outfalls from hydrocarbon processing plants. Roddie et al. (1994) found high levels of mortality of Corophium at sites contaminated with crude oil.

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

This pressure is Not assessed but any evidence is presented where available.

In general, crustaceans are widely reported to be intolerant of synthetic chemicals (Cole et al., 1999) and intolerance to some specific chemicals has been observed in amphipods. Species of a different genus are likely to differ in their susceptibility to synthetic chemicals and that this may be related to differences in their physiology (Powell, 1979). Corophium volutator is paralysed by pyrethrum based insecticide sprayed onto the surface of the mud (Gerdol & Hughes, 1993) and pyrethrum would probably cause significant mortalities if it found its way into estuaries from agricultural runoff. Nonylphenol is an anthropogenic pollutant that regularly occurs in water bodies, it is an oestrogen mimic that is produced during the sewage treatment of non-ionic surfactants and can affect Corophium volutator (Brown et al., 1999). Nonylphenol is a hydrophobic molecule and often becomes attached to sediment in water bodies. This will make nonylphenol available for ingestion by Corophium volutator in estuaries where much of the riverine water-borne sediment flocculates and precipitates out of suspension to form mudflats. Nonylphenol is not lethal to Corophium volutator but does reduce growth and has the effect of causing the secondary antennae of males to become enlarged, which can make the amphipods more vulnerable to predators (Brown et al., 1999). Corophium volutator is killed by 1% ethanol if exposed for 24 hours or more but can withstand higher concentrations in short pulses. Such short pulses, however, have the effect of rephasing the diel rhythm and will delay the timing of swimming activity for the duration of the ethanol pulse (Harris & Morgan, 1984b).

The anti-parasite compound ivermectin is highly toxic to benthic polychaetes and crustaceans (Black et al., 1997; Collier & Pinn, 1998; Grant & Briggs, 1998, cited in Wilding & Hughes, 2010). OSPAR (2000) stated that, at that time, ivermectin was not licensed for use in mariculture but was incorporated into the feed as a treatment against sea lice at some farms. Ivermectin has the potential to persist in sediments, particularly fine-grained sediments at sheltered sites. Data from a farm in Galway, Ireland indicated that ivermectin was detectable in sediments adjacent to the farm at concentrations up to 6.8 μm/kg and to a depth of 9 cm (reported in OSPAR, 2000). Infaunal polychaetes have been affected by deposition rates of 78-780 mg ivermectin/m2. Furthermore, Polydora ciliata was abundant at polluted sites close to acidified, halogenated effluent discharge from a bromide-extraction plant in Amlwch, Anglesey (Hoare & Hiscock, 1974). Spionid polychaetes were found by McLusky (1982) to be relatively resistant of distillation and petrochemical industrial waste in Scotland.

No evidence (NEv) Not relevant (NR) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Corophium volutator readily absorbs radionuclides such as americium and plutonium from water and contaminated sediments (Miramand et al., 1982). However, the effect of contamination of the individuals was not known but accumulation through the food chain was assumed (Miramand et al., 1982). There was 'No evidence' on which to base an assessment.

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

This pressure is Not assessed.

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

Riedel et al. (2012) assessed the response of benthic macrofauna to hypoxia advancing to anoxia in the Mediterranean. The hypoxic and anoxic conditions were created for 3-4 days in a box that enclosed in-situ sediments. Polychaetes appeared to be sensitive to hypoxia, as only 10% of polychaetes survived. In general, epifauna were more sensitive than infauna, mobile species more sensitive than sedentary species and predatory species more sensitive than suspension and deposit feeders. The test conditions did not lead to the production of hydrogen sulphide that may have reduced mortalities compared to other observations.

Amphipods appear not to be tolerant of reduced oxygenation. For example, Ampelisca agassizi was reported to be intolerant of hypoxia (Diaz & Rosenberg, 1995) and Jassa falcata, another tube building amphipod species, was absent from Californian harbours with low oxygen concentrations (0-2.5 mg/l).  Polydora ciliata is repeatedly found at localities with oxygen deficiency (Pearson & Rosenberg, 1978). For example, in polluted waters in Los Angeles and Long Beach harbours Polydora ciliata was present in the oxygen range 0.0-3.9 mg/l and the species was abundant in hypoxic fjord habitats (Rosenberg, 1977). Furthermore, in a study investigating a polychaete community in the north-west Black Sea, Polydora ciliata was observed in all four study sites, including those severely affected by eutrophication and hypoxia as a result of the discharge of wastewaters (Vorobyova et al., 2008). However, Polydora ciliata is unlikely to be able to resist anoxic conditions. Hansen et al. (2002) reported near total extinction of all metazoan in the Mariager Fjord (Denmark), including Polydora spp. after a severe hypoxia event that resulted in complete anoxia in the water column for two weeks. Additionally, Como & Magni (2009) investigated seasonal variations in benthic communities known to be affected by episodic events of hypoxia. The authors observed that the abundance of Polydora ciliata varied seasonally, decreasing during the summer months, and suggested it could be explained by the occurrence of hypoxic/anoxic conditions and sulphidic sediments during the summer. No details of the levels of dissolved oxygen leading to these community responses were provided. Other polychaetes in the biotope are also likely to be able to deal with hypoxia. For example, during low tide, the polychaete Scoloplos armiger survives deoxygenation by ascending into the oxidative layer where it can maintain aerobic metabolism. In laboratory conditions, Scoloplos armiger survived low oxygen conditions for 40 hours (Schöttler & Grieshaber, 1988).

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 (<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. 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.

Sensitivity assessment. Cole et al. (1999) suggested possible adverse effects on marine species exposed to dissolved oxygen concentrations below 4 mg/l and probable adverse effects below 2 mg/l. The characteristic polychaete Polydora ciliata is repeatedly found at localities with oxygen deficiency (Pearson & Rosenberg, 1978) and seems to only be affected by severe deoxygenation episodes. Other polychaetes in the biotope are likely to behave similarly. However, the mortality of tube building Ampelisca spp. species is likely. Therefore, resistance to deoxygenation at the pressure benchmark level is assessed as 'Low' but resilience is likely to be 'High so that biotope sensitivity is assessed as 'Low' sensitivity to exposure to a dissolved oxygen concentration of less than or equal to 2 mg/l for 1 week.

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

Connor et al. (2004) suggested that this biotope was associated with moderate organic enrichment. Amphipods appear to be tolerant of and indeed prefer, high nutrient levels. Diaz et al. (2008) reported the change in the abundance of Ampelisca tube mats in Boston Harbour, USA when wastewater outfalls were moved from onshore to offshore. Boston Harbour became changed from anaerobic to more aerobic between 1992 and 2006.  Ampelisca mats became widespread in 1992 and then declined by 2000. Diaz et al. (2008) suggested that the optimal organic loading for the maintenance of large areas of amphipod tube mats around 500 gC per square meter per year; above or below which levels, the mats declined. 

Polydora ciliata is often found in environments subject to high levels of nutrients. For example, the species was abundant in areas of the Firth of Forth exposed to high levels of sewage pollution (Smyth, 1968), in nutrient-rich sediments in the Mondego Estuary, Portugal (Pardal et al., 1993), and the coastal lagoon Lago Fusaro, Naples (Sordino et al., 1989). The extensive growths of Polydora ciliata in mat formations were recorded at West Ganton, in the Firth of Forth, prior to the introduction of the Sewage Scheme (Read et al., 1983). The abundance of the species was probably associated with their ability to use the increased availability of nutrients as a food source and silt for tube building.

Sensitivity assessment. This pressure relates to increased levels of nitrogen, phosphorus and silicon in the marine environment compared to background concentrations. The characterizing species of this biotope are likely to be able to resist and be favoured by nutrient enrichment where increased availability of nutrients may be used as a source of food (Hiscock et al., 2005a).  Nevertheless, the biotope is assessed as 'Not Sensitive' at the pressure benchmark level, which is set at compliance with Water Framework Directive (WFD) criteria for good status, based on nitrogen concentration (UKTAG, 2014).

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

Connor et al. (2004) suggested that this biotope was associated with moderate organic enrichment. Amphipods appear to be tolerant of and indeed prefer, high nutrient levels. Diaz et al. (2008) reported the change in the abundance of Ampelisca tube mats in Boston Harbour, USA when wastewater outfalls were moved from onshore to offshore. Boston Harbour became changed from anaerobic to more aerobic between 1992 and 2006.  Ampelisca mats became widespread in 1992 and then declined by 2000. Diaz et al. (2008) suggested that the optimal organic loading for the maintenance of large areas of amphipod tube mats around 500 gC per square meter per year; above or below which levels the mats declined.  Gittenberger & Van Loon (2011) assigned Ampelisca brevicornis to AMBI Group II "Species indifferent to enrichment, always present in low densities with non-significant variations with time (from initial state, to slight unbalance). These include suspension feeders, less selective carnivores and scavengers".

Polydora ciliata is often found in environments subject to high levels of nutrients. For example, the species was abundant in areas of the Firth of Forth exposed to high levels of sewage pollution (Smyth, 1968), in nutrient-rich sediments in the Mondego Estuary, Portugal (Pardal et al., 1993), and the coastal lagoon Lago Fusaro in Naples (Sordino et al., 1989). The extensive growths of Polydora ciliata in mat formations were recorded at West Ganton, in the Firth of Forth, prior to the introduction of the Sewage Scheme (Read et al., 1983). The abundance of the species was probably associated with their ability to use the increased availability of nutrients as a food source and silt for tube building. In colonization experiments in an organically polluted fjord receiving effluent discharge from Oslo, Polydora ciliata had settled in large numbers within the first month (Green, 1983). However, Callier et al. (2007) investigated the spatial distribution of macrobenthos under a suspended mussel culture, in eastern Canada, where the sedimentation of organic matter to the bottom was approx. 1-3 gC/m2/day. Polydora ciliata was recorded as absent in the sites under the suspended mussel farm after one year and as dominant in reference areas of the study.  Como & Magni (2009) investigated seasonal variations in benthic communities known to be affected by episodic events of sediment over-enrichment. The authors observed that abundance of Polydora ciliata varied seasonally, and suggested this could be a result major accumulation of organic carbon-binding fine sediments in the study site.  Studies by Almeda et al. (2009) and Pedersen et al. (2010) investigated larval energetic requirements for Polydora ciliata and suggested maximum growth rates were reached at food concentrations ranging from 1.4 to 2.5 μg C/ml depending on larval size, and energetic carbon requirements of 0.09 to 3.15 μg C l/d, respectively. Borja et al. (2000) and Gittenberger & Van Loon (2011) both assigned Polydora ciliata to their AMBI Group IV ‘second-order opportunistic species (slight to pronounced unbalanced situations)’, However, Polydora ciliata can also occur in organically poor areas (Pearson & Rosenberg, 1978).

Sensitivity assessment. The evidence presented suggests that the characterizing species of this biotope are likely to be stimulated by enrichment and only affected by excessive organic enrichment (above the benchmark level). Therefore, resistance and resilience are assessed as 'High', and the biotope is assessed as 'Not Sensitive' to organic enrichment involving deposition of 100 gC/m2/yr.  It should be noted that a decrease in organic enrichment may be detrimental as the abundance and extent of Ampelisca tube mats require moderate organic enrichment (see Diaz et al., 2008) but a reduction in organic enrichment is not addressed by this pressure. 

Physical Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
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

SS.SMu.ISaMu.AmpPlon is characterized by sandy mud substratum.  A change to a rock or artificial substratum would result in the loss of suitable habitat and the characterizing species, significantly altering the character of the biotope. The biotope would be lost and/or reclassified.

Sensitivity assessment. Resistance to the pressure is considered 'None', and resilience 'Very Low' based on the permanent loss of suitable substratum to support the community of the characterizing tube-building polychaete and amphipod species. Sensitivity has been assessed as '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

SS.SMu.ISaMu.AmpPlon is characterized by sandy muds only (Connor et al., 2004). A change in sediment type by one Folk Class (based on the Long, 2006 simplification) would result in an increase in the fraction of sand and gravel in the substratum.  Ampelisca tube mat communities are recorded from sandflats (e.g. Mills, 1967, 1969) and could potentially survive a change to a fine sand substratum. However, the biotope would probably transition into its sandy equivalent SS.SSa.IFiSa.TbAmPo and be reclassified. Mackenzie et al. (2006) reported that Ampelisca abdita did not occur on medium to coarse sand because they had difficulty using in to build their tubes.  A change to coarse sand or gravels would no longer support the characteristic species and the biotope would be lost and/or reclassified. 

Sensitivity assessment. Resistance to the pressure is considered 'None', and resilience 'Very Low' based on the permanent loss of suitable substratum to support the community of the characterizing species. Sensitivity has been assessed as 'High'.

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

Removal of the substratum to 30 cm would result in the loss of the tube mat created by Ampelisca spp., and its associated species e.g. Polydora ciliata tubes. Recovery of sediments will be site-specific and will be influenced by currents, wave action and sediment availability (Desprez, 2000). Except in areas of mobile sands, the process tends to be slow (Kenny & Rees, 1996; Desprez, 2000). Boyd et al. (2005) found that in a site subject to long-term extraction (25 years), extraction scars were still visible after six years and sediment characteristics were still altered in comparison with reference areas with ongoing effects on the biota. The strongest currents are unable to transport gravel. A further implication of the formation of these depressions is a local drop in current strength associated with the increased water depth, resulting in deposition of finer sediments than those of the surrounding substrate (Desprez, 2000).

Sensitivity assessment. Resistance is assessed as 'None' as the extraction of the sediment will remove the characterizing and associated species present. Resilience is assessed as 'Medium' (see resilience section) and sediments may need to recover (where exposed layers are different). Biotope sensitivity is, therefore, assessed as 'Medium'.

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

The tubes of the polychaetes and amphipods are bound only with mucous and are, therefore, likely to be damaged or removed by abrasion. The Ampelisca dominated tube mat sits in the surface of the sediment. Mills (1967, 1969) noted that the tubes of Ampelisca increased the surface complexity of the sandflat in Barnstable Harbour, Massachusetts. However, their feeding decreased the median grain size of the sediment, towards fine particulates, and resulted in instability. Mills (1969) stated that minor breaks in the mat of Ampelisca tubes were sufficient to cause large areas to wash out, especially on windy days in a rising or falling tide.  The tube mat would probably be removed easily by the passage of bottom gears.  Therefore, resistance to abrasion is assessed as 'Low'  but resilience is probably  'High' so that sensitivity to surface abrasion is assessed as 'Low'

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

Activities that penetrate below the surface are likely to tear up and remove a significant proportion of the tube building community that characterize this biotope. Bergman & Van Santbrink (2000) found that direct mortality of gammarid amphipods, following a single passage of a beam trawl (in silty sediments where penetration is greater) was 28%. Furthermore, stomach analysis of fish caught scavenging in the tracks of beam trawls found parts of Ampelisca spp. indicating that these had been damaged and exposed by the trawl (Kaiser & Spencer, 1994). Experiments in shallow, wave disturbed areas, using a toothed, clam dredge, found that deposit-feeding polychaetes were more impacted than carnivorous species. Dredging resulted in reductions of >90% of Spiophanes bombyx immediately post dredging compared with before impact samples and the population reduction persisting for 90 days (although results may be confounded by storm events within the monitoring period which caused sediment mobility). The passage of the dredge across the sediment floor will have killed or injured some organisms that will then be exposed to potential predators/scavengers (Frid et al., 2000; Veale et al., 2000) providing a food source to mobile scavengers including these species. Bergman & Hup (1992) carried out a pre and post-experimental investigation using a 12 m beam trawl. The area was trawled three times over two days and samples taken up to two weeks after trawling. Some benthic species showed a 10-65% reduction in density after trawling the area three times. There was a significant lower of density (40-60%) of polychaete worms, including Spiophanes bombyx.

The Ampelisca dominated tube mat sits in the surface of the sediment. Mills (1967, 1969) noted that the tubes of Ampelisca increased the surface complexity of the sandflat in Barnstable Harbour, Massachusetts. However, their feeding decreased the median grain size of the sediment, towards fine particulates, and resulted in instability. Mills (1969) stated that minor breaks in the mat of Ampelisca tubes were sufficient to cause large areas to wash out, especially on windy days in a rising or falling tide.  The tube mat would probably be removed easily by the passage of bottom gears. 

Sensitivity assessment. The evidence presented suggests that the tube mat community and associated infauna may suffer significant mortality (>75%) as a result of penetrative activities of the seabed. Biotope resistance is, therefore, assessed as 'None' and recovery is assessed as 'Medium so that sensitivity is assessed as 'Medium'.

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

The biotope is likely to occur in relatively turbid waters that allow sediment deposition to support the community of characterizing tube-building polychaetes and amphipods, and, therefore, the species in the biotope are likely to be adapted to turbid conditions. Amphipods are tolerant of high turbidity and gather suspended sediment for the construction of tubes. Mills (1967) reported that feeding by Ampelisca vadorum and Ampelisca abdita were initiated by the turbidity of the water surrounding the tubes. However, the feeding structures of suspension feeders such as Ampelisca spp. may become clogged by large increases in suspended sediment or feeding may be terminated, compromising growth. However, Mackenzie et al. (2006) noted that the Ampelisca mat modified the mud surface substantially. The tube mat stabilized the mud surface, minimised the resuspension of mud by strong currents and the surface of the sediment becomes covered by their faecal pellets that produce little turbidity when in the water column.  Mackenzie et al. (2006) also noted that suspension feeding by the dense population of Ampelisca spp. reduces turbidity locally by capturing silt in their faeces.

Tube-building polychaetes are likely to be tolerant of high turbidity as they normally inhabit waters with high levels of suspended sediment which they actively fix in the process of tube making. For example, in the Firth of Forth, Polydora ciliata formed extensive mats in areas that had an average of 68 mg/l suspended solids and a maximum of approximately 680 mg/l indicating the species can tolerate different levels of suspended solids (Read et al., 1982; Read et al., 1983). Daro & Polk (1973) reported that the success of Polydora is directly related to the quantities of muds of any origin carried along by rivers or coastal currents. Deposit feeders and tube builders rely on siltation of suspended sediment. A decrease in suspended sediment will reduce this supply and therefore may compromise growth and reproduction.  Spiophanes bombyx is found in estuarine regions which experience high levels of turbidity. Spiophanes bombyx is a surface deposit feeder and relies on a supply of nutrients at the sediment surface. An increase in turbidity, reducing light availability may reduce primary production by phytoplankton in the water column. Although productivity in the biotope is secondary, a reduction in primary production in the water column may result in reduced food supply to deposit and suspension feeders, which in turn may affect growth rates and fecundity.

Sensitivity assessment. An increase in suspended solids at the pressure benchmark level is unlikely to affect the characterizing species of this biotope. However, a decrease in the suspended matter in the biotope could result in limitation of material for tube building activities and the loss of suitable substratum for colonization by new recruits of Polydora ciliata, in particular. Therefore, resistance is assessed as 'Low' (loss of 25-75%) and resilience is 'High' so that sensitivity is assessed as 'Low' at the benchmark level. 

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

The tubes of Ampelisca spp. are probably several centimetres in length, depending on species, and extend above the sediment surface. For example, the tubes of Ampelisca abdita in Barnstable Harbour were about 3.5 cm in length a protruded 1 cm above the sediment surface (Mills, 1967). The tube mats occur in sedimentary, muddy environments so that they are probably adapted to steady rates of sedimentation and require sediment to continue to feed and build their tubes. However, no information on their ability to burrow or 'grow up' through deposited sediment was found. Ampelisca brevicornis was characterized by Gittenberger & Van Loon (2011) in their index of sedimentation tolerance as Group II "species sensitive to high sedimentation. They prefer to live in areas with some sedimentation, but don’t easily recover from strong fluctuations in sedimentation". 

A Polydora mud can be up to 50 cm thick, but the animals themselves occupy only the first few centimetres. They either elongate their tubes or have left them to rebuild close to the surface. Munari & Mistri (2014) investigated the spatio-temporal variation pattern of a benthic community following deposition of dredged material, at a maximum thickness of 30–40 cm. Polydora ciliata was amongst the first colonizers of the newly deposited sediments. The authors suggested that individuals may have migrated vertically through the deep layer of dredged sand. This was based on the results of Roberts et al. (1998) who suggested 15 cm as the maximum depth of overburden through which benthic infauna can successfully migrate. After one year, no adverse impact of sand disposal on the benthic fauna was detected on the study site (Munari & Mistri, 2014).  Spiophanes bombyx was Group IV species: ‘Although they are sensitive to strong fluctuations in sedimentation, their populations recover relatively quickly and even benefit. This causes their population sizes to increase significantly in areas after a strong fluctuation in sedimentation’ (Gittenberger & Van Loon, 2011). 

Sensitivity assessment. The Ampelisca tube mat is probably of a similar height to a deposit of 5 cm of fine sediment.  Polydora ciliata and other characterizing polychaetes are likely to resist or relocate after smothering by 5 cm of sediment. However, this ‘light’ deposition of fine sediment is may cause some mortality of Ampelisca spp.. based on Gittenberger & Van Loon (2011).  Therefore, resistance is assessed as 'Medium' albeit with 'Low' confidence.  Resilience is probably 'High' and biotope sensitivity is assessed as 'Low' sensitivity to a ‘light’ deposit of up to 5 cm of fine material in a single discrete event.

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

The tubes of Ampelisca spp. are probably several centimetres in length, depending on species, and extend above the sediment surface. For example, the tubes of Ampelisca abdita in Barnstable Harbour were about 3.5 cm in length a protruded 1 cm above the sediment surface (Mills, 1967). The tube mats occur in sedimentary, muddy environments so that they are probably adapted to steady rates of sedimentation and require sediment to continue to feed and build their tubes. However, no information on their ability to burrow or 'grow up' through deposited sediment was found. Ampelisca brevicornis was characterized by Gittenberger & Van Loon (2011) in their index of sedimentation tolerance as Group II "species sensitive to high sedimentation. They prefer to live in areas with some sedimentation, but don’t easily recover from strong fluctuations in sedimentation". 

A Polydora mud can be up to 50 cm thick, but the animals themselves occupy only the first few centimetres. They either elongate their tubes or have left them to rebuild close to the surface. Munari & Mistri (2014) investigated the spatio-temporal variation pattern of a benthic community following deposition of dredged material, at a maximum thickness of 30–40 cm. Polydora ciliata was amongst the first colonizers of the newly deposited sediments. The authors suggested that individuals may have migrated vertically through the deep layer of dredged sand. This was based on the results of Roberts et al. (1998) who suggested 15 cm as the maximum depth of overburden through which benthic infauna can successfully migrate. After one year, no adverse impact of sand disposal on the benthic fauna was detected on the study site (Munari & Mistri, 2014).  Spiophanes bombyx was Group IV species: ‘Although they are sensitive to strong fluctuations in sedimentation, their populations recover relatively quickly and even benefit. This causes their population sizes to increase significantly in areas after a strong fluctuation in sedimentation’ (Gittenberger & Van Loon, 2011). 

Sensitivity assessment. The Ampelisca tube mat is probably only a few centimetres in height and would be completely buried under a 30 cm deposit of sediment.  Polydora ciliata and other characterizing polychaetes are likely to resist or relocate after smothering by 30 cm of sediment. However, this ‘heavy’ deposition of fine sediment is may cause significant mortality of Ampelisca spp.. based on Gittenberger & Van Loon (2011).  Therefore, resistance is assessed as 'Low' albeit with 'Low' confidence.  Resilience is probably 'High' and biotope sensitivity is assessed as 'Low' sensitivity to a ‘heavy' deposit of up to 30 cm of fine material in a single discrete event.

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) Not relevant (NR) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

No evidence was available on which to assess this pressure. However, Arendse & Barendregt (1981) manipulated magnetic fields to alter the geomagnetic orientation of the talitrid amphipod Orchestia cavimana.

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

There is no evidence to suggest that any of the species that characterize the biotope respond to noise or vibration at the level of the benchmark, so the biotope is assessed as Not relevant.

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

SS.SMU.ISaMu.AmpPlon is a sublittoral biotope (Connor et al., 2004) and therefore not directly dependent on sunlight. Although, the characteristic species may respond to light orientation or shading they do not depend on light for feeding. Shading may decrease the abundance of benthic diatoms and euglenoids but the characteristic Ampelisca spp. are suspension-feeders capable of taking a range of organic particulates. The biotope is considered to have 'High' resistance and, by default, 'High' resilience and is, therefore, is assessed as 'Not Sensitive' to the introduction of light.

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 biotopes restricted to open waters.

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

Polydora ciliata exhibits shadow responses and withdraws its palps into its burrow, which is believed to be a defence against predation. The withdrawal of the palps interrupts feeding and possibly respiration, although the species also shows habituation of the response (Kinne, 1970).  Polydora is unlikely to be sensitive to visual disturbance caused by passing shipping but may respond to passing divers at close range.  Other characterizing polychaetes, such as Spiophanes bombyx, also inhabit a tube so its visual range is probably very limited.  No information on the visual responses of Ampelisca spp. was found. Nevertheless, visual disturbance (as defined by the benchmark) is probably 'Not relevant'

Biological Pressures

Use / to open/close text displayedResistanceResilienceSensitivity
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 important characterizing species in the biotope are not cultivated or likely to be translocated. This pressure is, therefore considered 'Not relevant'.

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

There is no evidence on the presence of non-indigenous species or impacts of non-indigenous species relevant to this biotope. This pressure is, therefore considered 'Not relevant'.

No evidence (NEv) Not relevant (NR) No evidence (NEv)
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR
Q: NR
A: NR
C: NR

Introduced organisms (especially parasites or pathogens) are a potential threat in all coastal ecosystems. However, no information was found on microbial pathogens affecting Polydora ciliata. Amphipods may be infected by a number of parasites or pathogens that alter population numbers through changes in host condition, growth, behaviour and reproduction (Green Extabe & Ford, 2014). For example, infection by acanthocephalan larvae may alter the behaviour and responses of gammarid amphipods (Bethel & Holmes, 1977). The amphipod Orchestia gammarellus is host to the parasitic protist Marteilia that has a feminizing effect on populations, with higher ratios of females and intersex males in infected, estuarine populations (Ginsburger-Vogel & Desportes, 1979). Studies conducted in the Baltic Sea suggested that increased parasitism by trematode species has a detrimental effect on local amphipods (Meissner & Bick, 1999; Mouritsen & Jensen, 1997; cited in Shim et al., 2013).  

Sensitivity assessment. There are no records of the biotope being affected by the introduction of microbial pathogens or parasites in the British Isles, and there is not enough evidence to assess sensitivity. 

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

None of the characterizing species is known to be targeted directly by commercial or recreational fisheries or harvesting. Therefore, this pressure is assessed as 'Not relevant'.

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

Direct, physical impacts are assessed through the abrasion and penetration of the seabed pressures, while this pressure considers the ecological or biological effects of by-catch. The characterizing species in this biotope are highly likely to be damaged or directly removed by static or mobile gears that are targeting other species (see abrasion and penetration pressures).

Sensitivity assessment. Loss of the characterizing species of this biotope is likely to occur as a result of unintentional removal. Removal of the characterizing species would result in the biotope being lost. Thus, the biotope is considered to have a resistance of 'Low' to this pressure.  However, resilience is probably  'High' so that sensitivity is assessed as 'Low'.

Bibliography

  1. Aberkali, H.B. & Trueman, E.R., 1985. Effects of environmental stress on marine bivalve molluscs. Advances in Marine Biology, 22, 101-198.

  2. Almeda, R., Pedersen, T.M., Jakobsen, H.H., Alcaraz, M., Calbet, A. & Hansen, B.W., 2009. Feeding and growth kinetics of the planktotrophic larvae of the spionid polychaete Polydora ciliata (Johnston). Journal of Experimental Marine Biology and Ecology, 382 (1), 61-68.

  3. Anger V., 1984. Reproduction in Pygospio-elegans Spionidae in relation to its geographical origin and to environmental conditions a preliminary report. Fischer, A. and H.-D. Pfannenstiel,  Fortschritte der Zoologie. pp. 45-52.

  4. Arendse, M.C. & Barendregt, A., 1981. Magnetic orientation in the semi-terrestrial amphipod, Orchestia cavimana, and its interrelationship with photo-orientation and water loss. Physiological Entomology, 6 (4), 333-342.

  5. Bailey-Brook, J.H., 1976. Habitats of tubicolous polychaetes from the Hawaiian Islands and Johnston Atoll. Pacific Science, 30, 69-81.

  6. Barnes, R.S.K. & Hughes, R.N., 1992. An introduction to marine ecology. Oxford: Blackwell Scientific Publications.

  7. Barnes, R.S.K., 1994. The brackish-water fauna of northwestern Europe. Cambridge: Cambridge University Press.

  8. Bat, L., Raffaelli, D. & Marr, I.L., 1998. The accumulation of copper, zinc and cadmium by the amphipod Corophium volutator (Pallas). Journal of Experimental Marine Biology and Ecology, 223, 167-184.

  9. 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.

  10. Bergman, M.J.N. & Hup, M., 1992. Direct effects of beam trawling on macro-fauna in a sandy sediment in the southern North Sea. ICES Journal of Marine Science, 49, 5-11.

  11. Bergman, M.J.N. & Van Santbrink, J.W., 2000b. Fishing mortality of populations of megafauna in sandy sediments. In The effects of fishing on non-target species and habitats (ed. M.J. Kaiser & S.J de Groot), 49-68. Oxford: Blackwell Science.

  12. Bethel, W.M. & Holmes, J.C., 1977. Increased vulnerability of amphipods to predation owing to altered behavior induced by larval acanthocephalans. Canadian Journal of Zoology55 (1), 110-115.

  13. Beukema, J. & Dekker, R., 2005. Decline of recruitment success in cockles and other bivalves in the Wadden Sea: possible role of climate change, predation on postlarvae and fisheries. Marine Ecology Progress Series, 287, 149-167.

  14. Beukema, J.J. & De Vlas, J., 1979. Population parameters of the lugworm, Arenicola marina, living on tidal flats in the Dutch Wadden Sea. Netherlands Journal of Sea Research, 13, 331-353.

  15. Beukema, J.J., 1995. Long-term effects of mechanical harvesting of lugworms Arenicola marina on the zoobenthic community of a tidal flat in the Wadden Sea. Netherlands Journal of Sea Research, 33, 219-227.

  16. Black, K.D., Fleming, S. Nickell, T.D. & Pereira, P.M.F. 1997. The effects of ivermectin, used to control sea lice on caged farmed salmonids, on infaunal polychaetes. ICES Journal of Marine Science, 54, 276-279.

  17. Blackstock, J. & Barnes, M., 1982. The Loch Eil project: biochemical composition of the polychaete, Glycera alba (Müller), from Loch Eil. Journal of Experimental Marine Biology and Ecology, 57 (1), 85-92.

  18. Blanchard, M., 1997. Spread of the slipper limpet Crepidula fornicata (L.1758) in Europe. Current state and consequences. Scientia Marina, 61, Supplement 9, 109-118.

  19. Bohn, K., Richardson, C.A. & Jenkins, S.R., 2015. The distribution of the invasive non-native gastropod Crepidula fornicata in the Milford Haven Waterway, its northernmost population along the west coast of Britain. Helgoland Marine Research, 69 (4), 313.

  20. Boon, J.P., Zantvoort, M.B., Govaert, M.J.M.A. & Duinker, J.C ., 1985. Organochlorines in benthic polychaetes (Nephtys spp.) and sediments from the southern North Sea. Identification of individual PCB components. Netherlands Journal of Sea Research, 19, 93-109.

  21. Booth, A. M., Hansen, B.H., Frenzel, M., Johnsen, H. & Altin, D., 2015. Uptake and toxicity of methylmethacrylate‐based nanoplastic particles in aquatic organisms. Environmental Toxicology and Chemistry, 9999, 1–9.

  22. Borja, A., Franco, J. & Perez, V., 2000. A marine biotic index to establish the ecological quality of soft-bottom benthos within European estuarine and coastal environments. Marine Pollution Bulletin, 40 (12), 1100-1114.

  23. Boulcott, P. & Howell, T.R.W., 2011. The impact of scallop dredging on rocky-reef substrata. Fisheries Research (Amsterdam), 110 (3), 415-420.

  24. Bousfield, E.L., 1973. Shallow-water gammaridean Amphipoda of New England. London: Cornell University Press.

  25. Boyd, S., Limpenny, D., Rees, H. & Cooper, K., 2005. The effects of marine sand and gravel extraction on the macrobenthos at a commercial dredging site (results 6 years post-dredging). ICES Journal of Marine Science: Journal du Conseil, 62 (2), 145-162.

  26. Bradshaw, C., Veale, L.O., Hill, A.S. & Brand, A.R., 2002. The role of scallop-dredge disturbance in long-term changes in Irish Sea benthic communities: a re-analysis of an historical dataset. Journal of Sea Research, 47, 161-184.

  27. Brils, J.M., Huwer, S.L., Kater, B.J., Schout, P.G., Harmsen, J., Delvigne, G.A.L. & Scholten, M.C.T., 2002. Oil effect in freshly spiked marine sediment on Vibrio fischeri, Corophium volutator, and Echinocardium caudatum. Environmental Toxicology and Chemistry, 21, 2242-2251.

  28. Brown, R.J., Conradi, M. & Depledge, M.H., 1999. Long-term exposure to 4-nonylphenol affects sexual differentiation and growth of the amphipod Corophium volutator (Pallas, 1766). Science of the Total Environment, 233, 77-88.

  29. Bryan, G.W. & Gibbs, P.E., 1983. Heavy metals from the Fal estuary, Cornwall: a study of long-term contamination by mining waste and its effects on estuarine organisms. Plymouth: Marine Biological Association of the United Kingdom. [Occasional Publication, no. 2.]

  30. 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.

  31. Bryant, V., McLusky, D.S., Roddie, K. & Newbery, D.M., 1984. Effect of temperature and salinity on the toxicity of chromium to three estuarine invertebrates (Corophium volutator, Macoma balthica, Nereis diversicolor). Marine Ecology Progress Series, 20, 137-149.

  32. Bryant, V., Newbery, D.M., McLusky, D.S. & Campbell, R., 1985. Effect of temperature and salinity on the toxicity of arsenic to three estuarine invertebrates (Corophium volutator, Macoma balthica, Tubifex costatus). Marine Ecology Progress Series, 24, 129-137.

  33. Bryant, V., Newbery, D.M., McLusky, D.S. & Campbell, R., 1985a. Effect of temperature and salinity on the toxicity of nickel and zinc to two estuarine invertebrates (Corophium volutator, Macoma balthica). Marine Ecology Progress Series, 24, 139-153.

  34. Callier, M. D., McKindsey, C.W. & Desrosiers, G., 2007. Multi-scale spatial variations in benthic sediment geochemistry and macrofaunal communities under a suspended mussel culture. Marine Ecology Progress Series, 348, 103-115.

  35. Capasso, E., Jenkins, S., Frost, M. & Hinz, H., 2010. Investigation of benthic community change over a century-wide scale in the western English Channel. Journal of the Marine Biological Association of the United Kingdom, 90 (06), 1161-1172.

  36. 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.], http://www.ukmarinesac.org.uk/

  37. Collie, J.S., Escanero, G.A. & Valentine, P.C., 1997. Effects of bottom fishing on the benthic megafauna of Georges Bank. Marine Ecology Progress Series, 155, 159-172.

  38. Collie, J.S., Hall, S.J., Kaiser, M.J. & Poiner, I.R., 2000. A quantitative analysis of fishing impacts on shelf-sea benthos. Journal of Animal Ecology, 69 (5), 785–798.

  39. 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.

  40. Como, S. & Magni, P., 2009. Temporal changes of a macrobenthic assemblage in harsh lagoon sediments. Estuarine, Coastal and Shelf Science, 83 (4), 638-646.

  41. 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.

  42. Connor, D.W., Allen, J.H., Golding, N., Howell, K.L., Lieberknecht, L.M., Northen, K.O. & Reker, J.B., 2004. The Marine Habitat Classification for Britain and Ireland. Version 04.05. ISBN 1 861 07561 8. In JNCC (2015), The Marine Habitat Classification for Britain and Ireland Version 15.03. [2019-07-24]. Joint Nature Conservation Committee, Peterborough. Available from https://mhc.jncc.gov.uk/

  43. 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.

  44. Conradi, M. & Depledge, M.H., 1999. Effects of zinc on the life-cycle, growth and reproduction of the marine amphipod Corophium volutator. Marine Ecology Progress Series, 176, 131-138.

  45. Cooper, K., Ware, S., Vanstaen, K. & Barry, J., 2011. Gravel seeding - A suitable technique for restoring the seabed following marine aggregate dredging? Estuarine, Coastal and Shelf Science, 91 (1), 121-132.

  46. Crisp, D.J. (ed.), 1964. The effects of the severe winter of 1962-63 on marine life in Britain. Journal of Animal Ecology, 33, 165-210.

  47. Cryer, M., Whittle, B.N. & Williams, K., 1987. The impact of bait collection by anglers on marine intertidal invertebrates. Biological Conservation, 42, 83-93.

  48. Daro, M.H. & Polk, P., 1973. The autecology of Polydora ciliata along the Belgian coast. Netherlands Journal of Sea Research, 6, 130-140.

  49. Dauvin, J-C. & Bellan-Santini, D., 1990. An overview of the amphipod genus Haploops (Ampeliscidae). Journal of the Marine Biological Association of the United Kingdom, 70, 887-903.

  50. Dauvin, J.C., 1988b. Biologie, dynamique, et production de populations de Crustacés amphipodes de la Manche occidentale. 1. Ampelisca tenuicornis Liljeborg. Journal of Experimental Marine Biology and Ecology, 118, 55-84.

  51. Dauvin, J.C., 1988c. The life cycle, population dynamics and production of the populations of amphipod crustaceans of the English Channel. 3. Ampelisca typica (Bate). Journal of Experimental Marine Biology and Ecology, 121, 1-22.

  52. Dauvin, J.C., 1988d. Life cycle, dynamics, and productivity of Crustacea-Amphipoda from the western English Channel. 4. Ampelisca armoricana Bellan-Santini et Dauvin. Journal of Experimental Marine Biology and Ecology, 123, 235-252

  53. Dauvin, J.C., 1988e. Biologie, dynamique, et production de populations de crustacés amphipodes de la Manche occidentale. 2. Ampelisca brevicornis (Costa). Journal of Experimental Marine Biology and Ecology, 119, 213-233.

  54. Dauvin, J.C., 1989. Life cycle, dynamics and productivity of Crustacea-Amphipoda from the western English Channel. 5. Ampelisca sarsi Chevreux. Journal of Experimental Marine Biology and Ecology, 128, 31-56.

  55. 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.

  56. 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.

  57. De Montaudouin, X. & Sauriau, P.G., 1999. The proliferating Gastropoda Crepidula fornicata may stimulate macrozoobenthic diversity. Journal of the Marine Biological Association of the United Kingdom, 79, 1069-1077.

  58. De-la-Ossa-Carretero, J., Del-Pilar-Ruso, Y., Loya-Fernández, A., Ferrero-Vicente, L., Marco-Méndez, C., Martinez-Garcia, E. & Sánchez-Lizaso, J., 2016. Response of amphipod assemblages to desalination brine discharge: impact and recovery. Estuarine, Coastal and Shelf Science, 172, 13-23

  59. Degraer, S., Wittoeck, J., Appeltans, W., Cooreman, K., Deprez, T., Hillewaert, H., Hostens, K., Mees, J., Vanden Berghe, E. & Vincx, M., 2006. The macrobenthos atlas of the Belgian part of the North Sea. Belgian Science Policy, Brussels.

  60. Desprez, M., 2000. Physical and biological impact of marine aggregate extraction along the French coast of the Eastern English Channel: short- and long-term post-dredging restoration. ICES Journal of Marine Science, 57 (5), 1428-1438.

  61. Diaz, R.J. & Rosenberg, R., 1995. Marine benthic hypoxia: a review of its ecological effects and the behavioural responses of benthic macrofauna. Oceanography and Marine Biology: an Annual Review, 33, 245-303.

  62. Diaz, R.J., Rhoads, D.C., Blake, J.A., Kropp, R.K. & Keay, K.E., 2008. Long-term trends of benthic habitats related to reduction in wastewater discharge to Boston Harbor. Estuaries and Coasts, 31 (6), 1184-1197. https:/doi.org/10.1007/s12237-008-9094-z
  63. Diaz-Castaneda, V., Richard, A. & Frontier, S., 1989. Preliminary results on colonization, recovery and succession in a polluted areas of the southern North Sea (Dunkerque's Harbour, France). Scientia Marina, 53, 705-716.

  64. Dittmann, S., Günther, C-P. & Schleier, U., 1999. Recolonization of tidal flats after disturbance. In The Wadden Sea ecosystem: stability, properties and mechanisms (ed. S. Dittmann), pp.175-192. Berlin: Springer-Verlag.

  65. Dorsett, D.A., 1961. The reproduction and maintenance of Polydora ciliata (Johnst.) at Whitstable. Journal of the Marine Biological Association of the United Kingdom, 41, 383-396.

  66. Drolet, D., Kennedy, K. & Barbeau, M.A., 2013. Winter population dynamics and survival strategies of the intertidal mudflat amphipod Corophium volutator (Pallas). Journal of Experimental Marine Biology and Ecology, 441, 126-137.

  67. Eagle, R.A., 1975. Natural fluctuations in a soft bottom benthic community. Journal of the Marine Biological Association of the United Kingdom, 55, 865-878.

  68. Eisler, R., 1977. Toxicity evaluation of a complex meta mixture to the softshell clam Mya arenaria. Marine Biology, 43, 265-276.

  69. 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.

  70. Emery, K.O., Stevenson, R.E., Hedgepeth, J.W., 1957. Estuaries and lagoons. In Treatise on marine ecology and paleoecology. vol. 1. Ecology, (ed. J.W. Hedgpeth), Geological Society of America, Memoir 67, pp. 673-750. Waverley Press, Baltimore, Mayland.

  71. Essink, K., 1999. Ecological effects of dumping of dredged sediments; options for management. Journal of Coastal Conservation, 5, 69-80.

  72. Fahy, E., Carroll, J. & O'Toole, M., 2003. A preliminary account of fisheries for the surf clam Spisula solida (L) (Mactracea) in Ireland [On-line] http://www.marine.ie, 2004-03-16

  73. 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.

  74. Fish, J.D. & Fish, S., 1974. The breeding cycle and growth of Hydrobia ulvae in the Dovey estuary. Journal of the Marine Biological Association of the United Kingdom, 54, 685-697.

  75. Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.

  76. Fish, J.D. & Mills, A., 1979. The reproductive biology of Corophium volutator and C. arenarium (Crustacea: Amphipoda). Journal of the Marine Biological Association of the United Kingdom, 59, 355-368.

  77. Flach, E.C., 1993. The distribution of the amphipod Corophium arenarium in the Dutch Wadden Sea- relationships with sediment composition and the presence of cockles and lugworms. Netherlands Journal of Sea Research, 31 (3), 281-290.

  78. Flach, E.C. & De Bruin, W., 1993. Effects of Arenicola marina and Cerastoderma edule on distribution, abundance and population structure of Corophium volutator in Gullmarsfjorden western Sweden. Sarsia, 78, 105-118.

  79. Flach, E.C. & De Bruin, W., 1994. Does the activity of cockles, Cerastoderma edule (L.) and lugworms, Arenicola marina (L.), make Corophium volutator Pallas more vulnerable to epibenthic predators: a case of interaction modification? Journal of Experimental Marine Biology and Ecology, 182, 265-285.

  80. Folk, R.L., 1954. The distinction between grain size and mineral composition in sedimentary-rock nomenclature. 62The Journal of Geology, 344-359.

  81. Forbes, M.R., Boates, S.J., McNeil, N.L. & Brison, A.E., 1996. Mate searching by males of the intertidal amphipod Corophium volutator (Pallas). Canadian Journal of Zoology, 74, 1479-1484.

  82. Ford, R.B. & Paterson, D.M., 2001. Behaviour of Corophium volutator in still versus flowing water. Estuarine, Coastal and Shelf Science, 52, 357-362.

  83. Fowler, S.L., 1999. Guidelines for managing the collection of bait and other shoreline animals within UK European marine sites. Natura 2000 report prepared by the Nature Conservation Bureau Ltd. for the UK Marine SACs Project, 132 pp., Peterborough: English Nature (UK Marine SACs Project)., http://www.english-nature.org.uk/uk-marine/reports/reports.htm

  84. Fretter, V. & Graham, A., 1981. The Prosobranch Molluscs of Britain and Denmark. Part 6. olluscs of Britain and Denmark. part 6. Journal of Molluscan Studies, Supplement 9, 309-313.

  85. Frid, C.L., Harwood, K.G., Hall, S.J. & Hall, J.A., 2000. Long-term changes in the benthic communities on North Sea fishing grounds. ICES Journal of Marine Science, 57 (5), 1303.

  86. Gamble, J., 1970. Anaerobic survival of the crustaceans Corophium volutatorC. arenarium and Tanais chevreuxiJournal of the Marine Biological Association of the United Kingdom50 (03), 657-671.

  87. Gamenick, I., Jahn, A., Vopel, K. & Giere, O., 1996. Hypoxia and sulphide as structuring factors in a macrozoobenthic community on the Baltic Sea shore: Colonization studies and tolerance experiments. Marine Ecology Progress Series, 144, 73-85.

  88. Gameson, 1982. The quality of the Humber Estuary, 1961-1981, Yorkshire Water Authority.

  89. Gerdol, V. & Hughes, R.G., 1993. Effect of the amphipod Corophium volutator on the colonisation of mud by the halophyte Salicornia europea. Marine Ecology Progress Series, 97, 61-69.

  90. Gibson, G.D. & Harvey, J., 2000. Morphogenesis during asexual reproduction in Pygospio elegans Claparede (Annelida, Polychaeta). The Biological Bulletin, 199 (1), 41-49.

  91. Gilkinson, K., Paulin, M., Hurley, S. & Schwinghamer, P., 1998. Impacts of trawl door scouring on infaunal bivalves: results of a physical trawl door model/dense sand interaction. Journal of Experimental Marine Biology and Ecology, 224 (2), 291-312.

  92. Gilkinson, K.D., Gordon, D.C., MacIsaac, K.G., McKeown, D.L., Kenchington, E.L., Bourbonnais, C. & Vass, W.P., 2005. Immediate impacts and recovery trajectories of macrofaunal communities following hydraulic clam dredging on Banquereau, eastern Canada. ICES Journal of Marine Science: Journal du Conseil, 62 (5), 925-947.

  93. Ginsburger-Vogel, T. & Desportes, I., 1979. Structure and biology of Marteilia sp. in the amphipod Orchestia gammarellus. Marine Fisheries Review, 41, 3-7.

  94. Gittenberger, A. & Van Loon, W.M.G.M., 2011. Common Marine Macrozoobenthos Species in the Netherlands, their Characterisitics and Sensitivities to Environmental Pressures. GiMaRIS report no 2011.08. DOI: 10.13140/RG.2.1.3135.7521

  95. Gogina, M., Glockzin. M. & Zettler, M.L., 2010. Distribution of benthic macrofaunal communities in the western Baltic Sea with regard to near-bottom environmental parameters. 2. Modelling and prediction. Journal of Marine Systems, 80, 57-70. 

  96. Grassle, J.F. & Grassle, J.P., 1974. Opportunistic life histories and genetic systems in marine benthic polychaetes. Journal of Marine Research, 32, 253-284.

  97. Gray, J.S., Clarke, K.R., Warwick, R.M. & Hobbs, G., 1990. Detection of initial effects of pollution on marine benthos - an example from the Ekofisk and Eldfisk oilfields, North Sea. Marine Ecology Progress Series, 66 (3), 285-299.

  98. Green, J., 1961. A biology of Crustacea. London: H.F. & G. Witherby Ltd. 180 pp.

  99. Green, N.W., 1983. Key colonisation strategies in a pollution-perturbed environment. In Fluctuations and Succession in Marine Ecosystems: Proceedings of the 17th European Symposium on Marine Biology, Brest, France, 27 September - 1st October 1982. Oceanologica Acta, 93-97.

  100. Gudmundsson, H., 1985. Life history patterns of polychaete species of the family spionidae. Journal of the Marine Biological Association of the United Kingdom, 65, 93-111.

  101. Gulliksen, B., 1977. Studies from the “UWL Helgoland” on the macrobenthic fauna of rocks and boulders in Lübeck Bay (western Baltic Sea). Helgolander Wissenschaftliche Meeresuntersuchungen, 30(1-4), 519-526.

  102. Hall, S.J., 1994. Physical disturbance and marine benthic communities: life in unconsolidated sediments. Oceanography and Marine Biology: an Annual Review, 32, 179-239.

  103. Hansen, B. W., Stenalt, E., Petersen, J.K. & Ellegaard, C., 2002. Invertebrate re-colonisation in Mariager Fjord (Denmark) after severe hypoxia. I. Zooplankton and settlement. Ophelia 56 (3), 197-213.

  104. Harms, J. & Anger, K., 1983. Seasonal, annual, and spatial variation in the development of hard bottom communities. Helgoländer Meeresuntersuchungen, 36, 137-150.

  105. Harris, G.J. & Morgan, E., 1984a. The effects of salinity changes on the endogenous circa-tidal rhythm of the amphipod Corophium volutator (Pallas). Marine Behaviour and Physiology, 10, 199-217.

  106. Harris, G.J. & Morgan, E., 1984b. The effects of ethanol, valinomycin and cycloheximide on the endogenous circa-tidal rhythm of the estuarine amphipod Corophium volutator (Pallas). Marine Behaviour and Physiology, 10, 219-233.

  107. Hayward, P.J. 1994. Animals of sandy shores. Slough, England: The Richmond Publishing Co. Ltd. [Naturalists' Handbook 21.]

  108. Hayward, P.J. & Ryland, J.S. (ed.) 1995b. Handbook of the marine fauna of North-West Europe. Oxford: Oxford University Press.

  109. Hiscock, K., 1983. Water movement. In Sublittoral ecology. The ecology of shallow sublittoral benthos (ed. R. Earll & D.G. Erwin), pp. 58-96. Oxford: Clarendon Press.

  110. Hiscock, K., Langmead, O., Warwick, R. & Smith, A., 2005a. Identification of seabed indicator species to support implementation of the EU Habitats and Water Framework Directives. Report to the Joint Nature Conservation Committee and the Environment Agency The Marine Biological Association, Plymouth, 77 pp.

  111. Hjulström, F., 1939. Transportation of detritus by moving water: Part 1. Transportation. Recent Marine Sediments, a Symposium (ed. P.D. Trask), pp. 5-31. Dover Publications, Inc.

  112. Hoare, R. & Hiscock, K., 1974. An ecological survey of the rocky coast adjacent to the effluent of a bromine extraction plant. Estuarine and Coastal Marine Science, 2 (4), 329-348.

  113. Holmström, W.F. & Morgan, E., 1983b. The effects of low temperature pulses in rephasing the endogenous activity rhythm of Corophium volutator (Pallas). Journal of the Marine Biological Association of the United Kingdom, 63, 851-860.

  114. 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.

  115. Hughes, R.G., 1988. Dispersal by benthic invertebrates: the in situ swimming behaviour of the amphipod Corophium volutator. Journal of the Marine Biological Association of the United Kingdom, 68, 565-579.

  116. Huthnance, J., 2010. Ocean Processes Feeder Report. London, DEFRA on behalf of the United Kingdom Marine Monitoring and Assessment Strategy (UKMMAS) Community.

  117. Jacobs, R.P.W.M., 1980. Effects of the Amoco Cadiz oil spill on the seagrass community at Roscoff with special reference to the benthic infauna. Marine Ecology Progress Series, 2, 207-212.

  118. Jensen, K.T. & Kristensen, L.D., 1990. A field experiment on competition between Corophium volutator (Pallas) and Corophium arenarium Crawford (Crustacea: Amphipoda): effects on survival, reproduction and recruitment. Journal of Experimental Marine Biology and Ecology, 137, 1-24.

  119. Jensen, K.T. & Mouritsen K.N., 1992. Mass mortality in two common soft bottom invertebrates, Hydrobia ulvae and Corophium volutator, the possible role of trematodes. Helgolander Meeresuntersuchungen, 46, 329-339.

  120. JNCC, 2015. The Marine Habitat Classification for Britain and Ireland Version 15.03. (20/05/2015). Available from https://mhc.jncc.gov.uk/

  121. JNCC (Joint Nature Conservation Committee), 1999. Marine Environment Resource Mapping And Information Database (MERMAID): Marine Nature Conservation Review Survey Database. [on-line] http://www.jncc.gov.uk/mermaid

  122. Kaiser, M.J. & Spencer, B.E., 1994. Fish scavenging behaviour in recently trawled areas. Marine Ecology Progress Series, 112 (1-2), 41-49.

  123. Kaschl, A. & Carballeira, A., 1999. Behavioural responses of Venerupis decussata (Linnaeus, 1758) and Venerupis pullastra (Montagu, 1803) to copper spiked marine sediments. Boletin. Instituto Espanol de Oceanografia, 15, 383-394.

  124. Kenny, A.J. & Rees, H.L., 1996. The effects of marine gravel extraction on the macrobenthos: results 2 years post-dredging. Marine Pollution Bulletin, 32 (8-9), 615-622.

  125. Kesaniemi, J.E., Geuverink, E. & Knott, K.E., 2012. Polymorphism in developmental mode and its effect on population genetic structure of a Spionid Polychaete, Pygospio elegans. Integrative and Comparative Biology, 52 (1), 181-196.

  126. Kinne, O. (ed.), 1970. Marine Ecology: A Comprehensive Treatise on Life in Oceans and Coastal Waters. Vol. 1 Environmental Factors Part 1. Chichester: John Wiley & Sons

  127. Kirby, R.R., Beaugrand, G. & Lindley, J.A., 2008. Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea. Limnology and Oceanography, 53 (5), 1805.

  128. Klawe, W.L. & Dickie, L.M., 1957. Biology of the bloodworm, Glycera dibranchiata Ehlers, and its relation to the bloodworm fishery of the Maritime Provinces. Bulletin of Fisheries Research Board of Canada, 115, 1-37.

  129. Kröncke, I., Dippner, J., Heyen, H. & Zeiss, B., 1998. Long-term changes in macrofaunal communities off Norderney (East Frisia, Germany) in relation to climate variability. Marine Ecology Progress Series, 167, 25-36.

  130. Kruse, I., Strasser, M. & Thiermann, F., 2004. The role of ecological divergence in speciation between intertidal and subtidal Scoloplos armiger (Polychaeta, Orbiniidae). Journal of Sea Research, 51, 53-62.

  131. Lagadeuc, Y., 1991. Mud substrate produced by Polydora ciliata (Johnston, 1828) (Polychaeta, Annelida) - origin and influence on fixation of larvae. Cahiers de Biologie Marine, 32, 439-450.

  132. Le Bot, S., Lafite, R., Fournier, M., Baltzer, A. & Desprez, M., 2010. Morphological and sedimentary impacts and recovery on a mixed sandy to pebbly seabed exposed to marine aggregate extraction (Eastern English Channel, France). Estuarine, Coastal and Shelf Science89221-233.

  133. Le Bris, H. & Glemarec, M., 1995. Macrobenthic communities of oxygen under-saturated ecosystems: The Bay of Vilaine, southern Brittany. Oceanologica Acta, 18, 573-581.

  134. Levell, D., Rostron, D. & Dixon, I.M.T., 1989. Sediment macrobenthic communities from oil ports to offshore oilfields. In Ecological Impacts of the Oil Industry, Ed. B. Dicks. Chicester: John Wiley & Sons Ltd.

  135. Long, D., 2006. BGS detailed explanation of seabed sediment modified Folk classification. Available from: http://www.emodnet-seabedhabitats.eu/PDF/GMHM3_Detailed_explanation_of_seabed_sediment_classification.pdf

  136. Lopez-Flores I., De la Herran, R., Garrido-Ramos, M.A., Navas, J.I., Ruiz-Rejon, C. & Ruiz-Rejon, M., 2004. The molecular diagnosis of Marteilia refringens and differentiation between Marteilia strains infecting oysters and mussels based on the rDNA IGS sequence. Parasitology19 (4), 411-419.

  137. Mackenzie, C.L., Pikanowski, R. & Mcmillan, D.G., 2006. Ampelisca amphipod tube mats may enhance abundance of Northern Quahogs Mercenaria mercenaria in muddy sediments. Journal of Shellfish Research, 25 (3), 841-847. https://doi.org/10.2983/0730-8000(2006)25[841:AATMME]2.0.CO;2
  138. Maurer, D. & Lethem, W., 1980. Dominant species of polychaetous annelids of Georges Bank. Marine Ecology Progress Series, 3, 135-144.

  139. Maurer, D., Keck, R.T., Tinsman, J.C. & Leathem, W.A., 1982. Vertical migration and mortality of benthos in dredged material: Part III—polychaeta. Marine Environmental Research, 6 (1), 49-68.

  140. McCabe, G.T. Jr., Hinton, S.A. & Emmett, R.L., 1998. Benthic invertebrates and sediment characteristics in a shallow navigation channel of the lower Columbia River. Northwest Science, 72, 116-126.

  141. McCall, P.L., 1977. Community patterns and adaptive strategies of the infaunal benthos of Long Island Sound. Journal of Marine Research, 35, 221-266.

  142. McCurdy, D.G., Boates, J.S. & Forbes, M.R., 2000. Reproductive synchrony in the intertidal amphipod Corophium volutator. Oikos, 88, 301-308.

  143. McDermott, J.J., 1984. The feeding biology of Nipponnemertes pulcher (Johnston) (Hoplonemertea), with some ecological implications. Ophelia, 23, 1-21.

  144. McLusky, D., Anderson, F. & Wolfe-Murphy, S., 1983. Distribution and population recovery of Arenicola marina and other benthic fauna after bait digging. Marine Ecology Progress Series, 11 (2), 173-179.

  145. McLusky, D.S., 1967. Some effects of salinity on the survival, moulting, and growth of Corophium volutator (Amphipoda). Journal of the Marine Biological Association of the United Kingdom, 47, 607-617.

  146. McLusky, D.S., 1968. Some effects of salinity on the distribution and abundance of Corophium volutator in the Ythan estuary. Journal of the Marine Biological Association of the United Kingdom, 48, 443-454.

  147. McLusky, D.S., 1982. The impact of petrochemical effluent on the fauna of an intertidal estuarine mudflat. Estuarine, Coastal and Shelf Science, 14, 489-499.

  148. 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.

  149. Meador, J.P., Varanasi, U. & Krone, C.A., 1993. Differential sensitivity of marine infaunal amphipods to tributyltin. Marine Biology, 116, 231-239.

  150. Meadows, P., 1964. Substrate selection by Corophium species: the particle size of substrates. The Journal of Animal Ecology, 33, 387-394.

  151. Meadows, P. & Reid, A., 1966. The behaviour of Corophium volutator (Crustacea: Amphipoda). Journal of Zoology 150(4): 387-399

  152. Meadows, P.S. & Ruagh, A.A., 1981. Temperature preferences and activity of Corophium volutator (Pallas) in a new choice apparatus. Sarsia, 66, 67-72.

  153. Meißner, K., Darr, A. & Rachor, E., 2008. Development of habitat models for Nephtys species (Polychaeta: Nephtyidae) in the German Bight (North Sea). Journal of Sea Research, 60 (4), 276-291.

  154. Marine Ecological Surveys Limited (MES), 2008. Marine Macrofauna Genus Trait Handbook. Marine Ecological Surveys Limited: Bath. 

  155. MES, 2010. Marine Macrofauna Genus Trait Handbook. Marine Ecological Surveys Limited. http://www.genustraithandbook.org.uk/

  156. Mills, A. & Fish, J., 1980. Effects of salinity and temperature on Corophium volutator and C. arenarium (Crustacea: Amphipoda), with particular reference to distribution. Marine Biology, 58 (2), 153-161.

  157. Mills, E.L., 1967. The biology of an ampeliscid amphipod crustacean sibling species pair. Journal of the Fisheries Research Board of Canada, 24, 305-355.

  158. Mills, E.L., 1969. The community concept in marine zoology, with comments on continua and instability in some marine communities: a review. Journal of the Fisheries Research Board of Canada, 26 (6), 1415-1428. DOI https://doi.org/10.1139/f69-132
  159. Miramand, P., Germain, P. & Camus, H., 1982. Uptake of americium and plutonium from contaminated sediments by three benthic species: Arenicola marina, Corophium volutator and Scrobicularia plana. Marine Ecology Progress Series, 7, 59-65.

  160. Moulaert, I. & Hostens, K., 2007. Post-extraction evolution of a macrobenthic community on the intensively extracted Kwintebank site in the Belgian part of the North Sea. CM Documents-ICES, (A:12).

  161. Mouritsen, K. N., Mouritsen, L.T. & Jensen, K.T., 1998. Change of topography and sediment characteristics on an intertidal mud-flat following mass-mortality of the amphipod Corophium volutator. Journal of the Marine Biological Association of the United Kingdom, 78 (4), 1167-1180.

  162. Mouritsen, K.N., Tompkins, D.M. & Poulin, R., 2005. Climate warming may cause a parasite-induced collapse in coastal amphipod populations. Oecologia, 146, 476-483.

  163. Munari, C. & Mistri, M., 2014. Spatio-temporal pattern of community development in dredged material used for habitat enhancement: A study case in a brackish lagoon. Marine Pollution Bulletin 89 (1–2), 340-347.

  164. Murina, V., 1997. Pelagic larvae of Black Sea Polychaeta. Bulletin of Marine Science, 60, 427-432.

  165. Mustaquim, J., 1986. Morphological variation in Polydora ciliata complex (Polychaeta, Annelida). Zoological Journal of the Linnean Society, 86, 75-88.

  166. Neal, K.J. & Avant, P. 2006. Corophium volutator A mud shrimp. 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://192.171.193.68/species/detail/1661

  167. Neves de Carvalho, A., Vaz, A.S.L., Sérgio, T.I.B. & Santos, P.J.T.d., 2013. Sustainability of bait fishing harvesting in estuarine ecosystems: Case study in the Local Natural Reserve of Douro Estuary, Portugal estuarinos: Caso de estudo na Reserva Natural Local do Estuário do Douro, Portugal. Revista de Gestão Costeira Integrada, 13 (2), 157-168.

  168. Newell, R., Seiderer, L. & Hitchcock, D., 1998. The impact of dredging works in coastal waters: a review of the sensitivity to disturbance and subsequent recovery of biological resources on the sea bed. Oceanography and Marine Biology: An Annual Review, 36, 127-178.

  169. Nichols, F.H. & Thompson, J.K., 1985. Persistence of an introduced mudflat community in South San Francisco Bay, California. Marine Ecology Progress Series24, 83-97.

  170. 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.

  171. OBIS, 2016. Ocean Biogeographic Information System (OBIS). http://www.iobis.org, 2016-03-15

  172. 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.

  173. OSPAR, 2000. OSPAR decision 2000/3 on the use of organic-phase drilling fluids (OPF) and the discharge of OPF-contaminated cuttings. Summary Record OSPAR 2000. OSPAR 00/20/1-E, Annex 18. Copenhagen, 26–30 June.

  174. Pardal, M.A., Marques, J.-C. & Bellan, G., 1993. Spatial distribution and seasonal variation of subtidal polychaete populations in the Mondego estuary (western Portugal). Cahiers de Biologie Marine, 34, 497-512.

  175. 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.

  176. Pedersen, T. M., Almeda, R., Fotel, F.L., Jakobsen, Hans H., Mariani, P. & Hansen, B.W., 2010. Larval growth in the dominant polychaete Polydora ciliata is food-limited in a eutrophic Danish estuary (Isefjord). Marine Ecology Progress Series, 407, 99-110.

  177. Petersen, C.G.J., 1918. The sea bottom and its production of fish food. A survey of the work done in connection with valuation of the Denmark waters from 1883-1917. Report of the Danish Biological Station, 25, 1-62.

  178. Picton, B.E. & Costello, M.J., 1998. BioMar biotope viewer: a guide to marine habitats, fauna and flora of Britain and Ireland. [CD-ROM] Environmental Sciences Unit, Trinity College, Dublin.

  179. 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.

  180. Poulin, R. & Mouritsen, K.N., 2006. Climate change, parasitism and the structure of intertidal ecosystems. Journal of Helminthology, 80 (2), 183-192.

  181. Powell, C.E., 1979. Isopods other than cyathura (Arthropoda: Crustacea: Isopoda). In Pollution ecology of estuarine invertebrates (ed. C.W. Hart & S.L.H. Fuller), 325-338. New York: Academic Press.

  182. Powilleit, M., Graf, G., Kleine, J., Riethmuller, R., Stockmann, K., Wetzel, M.A. & Koop, J.H.E., 2009. Experiments on the survival of six brackish macro-invertebrates from the Baltic Sea after dredged spoil coverage and its implications for the field. Journal of Marine Systems, 75 (3-4), 441-451.

  183. Raffaelli, D., Limia, J., Hull, S. & Pont, S., 1991. Interactions between the amphipod Corophium volutator and macroalgal mats on estuarine mudflats. Journal of the Marine Biological Association of the United Kingdom, 71, 899-908.

  184. Read, P.A., Anderson, K.J., Matthews, J.E., Watson, P.G., Halliday, M.C. & Shiells, G.M., 1982. Water quality in the Firth of Forth. Marine Pollution Bulletin, 13, 421-425.

  185. Read, P.A., Anderson, K.J., Matthews, J.E., Watson, P.G., Halliday, M.C. & Shiells, G.M., 1983. Effects of pollution on the benthos of the Firth of Forth. Marine Pollution Bulletin, 14, 12-16.

  186. Rees, E., Nicolaidou, A. & Laskaridou, P., 1976. The effects of storms on the dynamics of shallow water benthic associations. In Proceedings of the 11th European Symposium on Marine Biology, Galway, 5-11 October, 1976. Biology of benthic organisms (ed. B.F., Keegan; P., O'Ceidigh & P.J.S., Boaden), pp. 465-474.

  187. Reish, D.J., 1979. Bristle Worms (Annelida: Polychaeta) In Pollution Ecology of Estuarine Invertebrates, (eds. Hart, C.W. & Fuller, S.L.H.), 78-118. Academic Press Inc, New York.

  188. 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.

  189. Riedel, B., Zuschin, M. & Stachowitsch, M., 2012. Tolerance of benthic macrofauna to hypoxia and anoxia in shallow coastal seas: a realistic scenario. Marine Ecology Progress Series, 458, 39-52.

  190. Riera, R., Tuya, F., Ramos, E., Rodríguez, M. & Monterroso, Ó., 2012. Variability of macrofaunal assemblages on the surroundings of a brine disposal. Desalination, 291, 94-100.

  191. Roberts, R. D., Gregory, M.R. & Foster, B.A., 1998. Developing an efficient macrofauna monitoring index from an impact study—a dredge spoil example. Marine Pollution Bulletin, 36 (3), 231-235.

  192. Roche, C., Lyons, D.O.,O'Connor, B. 2007. Benthic surveys of sandbanks in the Irish Sea. Irish Wildlife Manuals, No. 29. National Parks and Wildlife Service, Department of Environment, Heritage and Local Government, Dublin, Ireland.

  193. Roddie, B., Kedwards, T., Ashby-Crane, R. & Crane, M., 1994. The toxicity to Corophium volutator (Pallas) of beach sand contaminated by a spillage of crude oil. Chemosphere, 29 (4), 719-727.

  194. Rosenberg, R., 1977. Benthic macrofaunal dynamics, production, and dispersion in an oxygen-deficient estuary of west Sweden. Journal of Experimental Marine Biology and Ecology, 26, 107-33.

  195. Sanders, H.L., 1978. Florida oil spill impact on the Buzzards Bay benthic fauna: West Falmouth. Journal of the Fisheries Board of Canada, 35 (5), 717-730.

  196. Sardá, R., Pinedo, S. & Martin, D., 1999. Seasonal dynamics of macroinfaunal key species inhabiting shallow soft-bottoms in the Bay of Blanes (NW Mediterranean). Publications Elsevier: Paris.

  197. Sardá, R., Pinedo, S., Gremare, A. & Taboada, S., 2000. Changes in the dynamics of shallow sandy-bottom assemblages due to sand extraction in the Catalan Western Mediterranean Sea. ICES Journal of Marine Science, 57 (5), 1446-1453.

  198. Schottler, U. & Grieshaber, M., 1988. Adaptation of the polychaete worm Scoloplos armiger to hypoxic conditions. Marine Biology, 99 (2), 215-222.

  199. SEEEC (Sea Empress Environmental Evaluation Committee), 1998. The environmental impact of the Sea Empress oil spill. Final Report of the Sea Empress Environmental Evaluation Committee, 135 pp., London: HMSO.

  200. Shepherd, P.C.F. & Boates, S.J., 1999. Effects of commercial baitworm harvest on semipalmated sandpipers and their prey in the Bay of Fundy hemispheric shorebird reserve. Conservation Biology, 13, 347-356.

  201. Shim, K.C., Koprivnikar, J. & Forbes, M.R., 2013. Variable effects of increased temperature on a trematode parasite and its intertidal hosts. Journal of Experimental Marine Biology and Ecology, 439, 61-68.

  202. Sinderman, C.J., 1990. Principle diseases of marine fish and shellfish, 2nd edition, Volume 2. Diseases of marine shellfish. Academic Press, 521 pp.

  203. Smyth, J.C., 1968. The fauna of a polluted site in the Firth of Forth. Helgolander Wissenschaftliche Meeresuntersuchungen, 17, 216-233.

  204. Snelgrove, P.V., Grassle, J.P., Grassle, J.F., Petrecca, R.F. & Ma, H., 1999. In situ habitat selection by settling larvae of marine soft‐sediment invertebrates. Limnology and Oceanography, 44 (5), 1341-1347.

  205. Sohtome, T., Wada, T., Mizuno, T., Nemoto, Y., Igarashi, S., Nishimune, A., Aono, T., Ito, Y., Kanda, J. & Ishimaru, T., 2014. Radiological impact of TEPCO's Fukushima Dai-ichi Nuclear Power Plant accident on invertebrates in the coastal benthic food web. Journal of Environmental Radioactivity, 138, 106-115.

  206. Sordino, P., Gambi, M.C. & Carrada, G.C., 1989. Spatio-temporal distribution of polychaetes in an Italian coastal lagoon (Lago Fusaro, Naples). Cahiers de Biologie Marine, 30, 375-391.

  207. Suchanek, T.H., 1993. Oil impacts on marine invertebrate populations and communities. American Zoologist, 33, 510-523.

  208. Sundborg, Å., 1956. The River Klarälven: a study of fluvial processes. Geografiska Annaler, 38 (2), 125-237.

  209. Thomas, R., 1975. Functional morphology, ecology, and evolutionary conservatism in the Glycymerididae (Bivalvia). Palaeontology, 18 (2), 217-254.

  210. Thorson, G., 1946. Reproduction and larval development of Danish marine bottom invertebrates, with special reference to the planktonic larvae in the Sound (Øresund). Meddelelser fra Kommissionen for Danmarks Fiskeri- Og Havundersögelser, Serie: Plankton, 4, 1-523.

  211. Thorson, G., 1957. Bottom communities (sublittoral or shallow shelf). Memoirs of the Geological Society of America, 67, 461-534.

  212. 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.

  213. Tuck, I.D., Hall, S.J., Robertson, M.R., Armstrong, E. & Basford, D.J., 1998. Effects of physical trawling disturbance in a previously unfished sheltered Scottish sea loch. Marine Ecology Progress Series, 162, 227-242.

  214. Ugolini, A., Ungherese, G., Somigli, S., Galanti, G., Baroni, D., Borghini, F., Cipriani, N., Nebbiai, M., Passaponti, M. & Focardi, S., 2008. The amphipod Talitrus saltator as a bioindicator of human trampling on sandy beaches. Marine Environmental Research, 65 (4), 349-357.

  215. UKTAG, 2014. UK Technical Advisory Group on the Water Framework Directive [online]. Available from: http://www.wfduk.org

  216. Valentine, P.C., Carman, M.R., Blackwood, D.S. & Heffron, E.J., 2007. Ecological observations on the colonial ascidian Didemnum sp. in a New England tide pool habitat. Journal of Experimental Marine Biology and Ecology, 342 (1), 109-121.

  217. Van Colen, C., Montserrat, F., Vincx, M., Herman, P.M.J., Ysebaert, T. & Degraer, S., 2010. Long-term divergent tidal flat benthic community recovery following hypoxia-induced mortality. Marine Pollution Bulletin 60 (2), 178-186.

  218. Van Dalfsen, J.A., Essink, K., Toxvig Madsen, H., Birklund, J., Romero, J. & Manzanera, M., 2000. Differential response of macrozoobenthos to marine sand extraction in the North Sea and the Western Mediterranean. ICES Journal of Marine Science, 57 (5), 1439-1445.

  219. Veale, L.O., Hill, A.S., Hawkins, S.J. & Brand, A.R., 2000. Effects of long term physical disturbance by scallop fishing on subtidal epifaunal assemblages and habitats. Marine Biology, 137, 325-337.

  220. Vorobyova, L., Bondarenko, O. & Izaak, O., 2008. Meiobenthic polychaetes in the northwestern Black Sea. Oceanological and Hydrobiological Studies, 37 (1), 43-55.

  221. Watkin, E.E., 1941. The yearly life cycle of the amphipod, Corophium volutator. The Journal of Animal Ecology, 10, 77-93.

  222. Watson, G.J., Farrell, P., Stanton, S. & Skidmore, L.C., 2007. Effects of bait collection on Nereis virens populations and macrofaunal communities in the Solent, UK. Journal of the Marine Biological Association of the United Kingdom, 87 (3), 703-716.

  223. Widdows, J., Bayne, B.L., Livingstone, D.R., Newell, R.I.E. & Donkin, P., 1979. Physiological and biochemical responses of bivalve molluscs to exposure to air. Comparative Biochemistry and Physiology, 62A, 301-308.

  224. Wilding T. & Hughes D., 2010. A review and assessment of the effects of marine fish farm discharges on Biodiversity Action Plan habitats. Scottish Association for Marine Science, Scottish Aquaculture Research Forum (SARF).

  225. Wilson, W.H. & Parker, K., 1996. The life history of the amphipod, Corophium volutator: the effects of temperature and shorebird predation. Journal of Experimental Marine Biology and Ecology, 196, 239-250.

  226. Ysebaert, T., Meire, P., Maes, D. & Buijs, J., 1993. The benthic macrofauna along the estuarine gradient of the Schelde estuary. Netherlands Journal of Aquatic Ecology, 27 (2-4), 327-341.

Citation

This review can be cited as:

Tyler-Walters, H. & De-Bastos, E., 2020. [Ampelisca] spp., [Photis longicaudata] and other tube-building amphipods and polychaetes in infralittoral sandy mud. 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. [cited 21-10-2020]. Available from: https://www.marlin.ac.uk/habitat/detail/1230

Last Updated: 20/03/2020