Semi-permanent tube-building amphipods and polychaetes in sublittoral sand

20-03-2002
Researched byWill Rayment Refereed byProf. Jean-Claude Dauvin
EUNIS CodeA5.234 EUNIS NameSemi-permanent tube-building amphipods and polychaetes in sublittoral sand

Summary

UK and Ireland classification

EUNIS 2008A5.234Semi-permanent tube-building amphipods and polychaetes in sublittoral sand
EUNIS 2006A5.234Semi-permanent tube-building amphipods and polychaetes in sublittoral sand
JNCC 2004SS.SSa.IFiSa.TbAmPoSemi-permanent tube-building amphipods and polychaetes in sublittoral sand
1997 BiotopeSS.IMU.MarMu.TubeAPSemi-permanent tube-building amphipods and polychaetes in sublittoral mud or muddy sand

Description

Sublittoral stable mud and muddy sands occurring over a wide depth range may support large populations of semi-permanent tube-building amphipods and polychaetes. This community is poorly known and appears to occur in restricted patches. Amphipods such as Ampelisca spp., Corophium spp. and Haploops tubicola have been described as occurring in high densities in such habitats (see Petersen, 1918; Thorson, 1957) and polychaetes such as Spiophanes bombyx and Polydora ciliata may also be conspicuously numerous. It may be that this community develops as a result of moderate nutrient enrichment. It is possible that this biotope may contain more than one entity as all the characterizing species listed need not occur simultaneously. (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Recorded distribution in Britain and Ireland

Common in lagoons on Shetland and Orkney. No records on mainland coasts of Britain or Ireland.

Depth range

-

Additional information

None

Listed By

Further information sources

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JNCC

Habitat review

Ecology

Ecological and functional relationships

  • The biotope is characterized by tube-building polychaetes and amphipods, with errant polychaetes and nemerteans foraging in the surrounding and underlying sediment.
  • The dominant tube-builders are the deposit feeding polychaetes Polydora ciliata, Spiophanes bombyx and Pygospio elegans. In areas of mud, the tubes built by Polydora ciliata can agglomerate and form layers of mud an average of 20 cm thick, occasionally up to 50 cm (Daro & Polk, 1973). The tube-building, suspension feeding amphipods Ampelisca sp. are present where the biotope occurs in shallow warm waters, while they are probably replaced by the very similar Haploops tubicola in deeper, colder waters (Dauvin & Bellan-Santini, 1990).
  • The feeding activities of high densities of Polydora ciliata may inhibit the establishment of other benthic species by removing settling and developing larvae (Daro & Polk, 1973).
  • Infaunal deposit feeding polychaetes include the burrow dwelling Arenicola marina, the sedentary Chaetozone setosa, the mobile detritivore Scoloplos armiger and species tolerant of nutrient enrichment including Capitomastus minimus and Capitella sp.
  • The amphipods and the infaunal annelid species in the biotope probably interfere strongly with each other. Adult worms probably reduce amphipod numbers by disturbing their burrows and tubes, while high densities of amphipods can prevent establishment of worms by consuming larvae and juveniles (Olafsson & Persson, 1986).
  • The biotope contains a number of infaunal bivalve species, including Abra alba, Fabulina fabula and Mysella bidentata, which probably both deposit feed and suspension feed depending on local environmental conditions.
  • Spatial competition probably occurs between the infaunal suspension feeders and deposit feeders. Reworking of sediment by deposit feeders, e.g. Arenicola marina, makes the substratum less stable, increases the suspended sediment and makes the environment less suitable for suspension feeders (Rhoads & Young, 1970). Tube building by amphipods stabilizes the sediment and arrests the shift towards a community consisting entirely of deposit feeders.
  • Amphipods are predated chiefly by nemertean worms. For example, the nemertean Nipponnemertes pulcher is the dominant predator in the Haploops community in the Danish Oeresund (McDermott, 1984).
  • Mobile, carnivorous polychaetes, including Anaitides mucosa, Eteone longa, Nephtys hombergi and Pholoe inornata, predate the smaller annelids and crustaceans.

Seasonal and longer term change

Temporal changes are likely to occur in the community due to seasonal recruitment processes. For example, the early reproductive period of Polydora ciliata often enables the species to be the first to colonize available substrata (Green, 1983). The settling of the first generation in April is followed by the accumulation and active fixing of mud continuously up to a peak during the month of May. The following generations do not produce a heavy settlement due to interspecific competition and heavy mortality of the larvae (Daro & Polk, 1973). Later in the year, the surface layer cannot hold the lower layers of the mud mat in place and they may be swept away by water currents. The substratum may now be colonized by the abundant larvae of other species in the water column.
There is a seasonal variation in planktonic production in surface waters which probably affects the food supply of the benthos in the biotope. Increased production by phytoplankton in spring and summer due to increased temperatures and irradiance is followed by phytoplankton sedimentation events which are correlated with seasonal variations in the organic content of benthic sediments (Thouzeau et al., 1996). These variations directly influence the food supply of the deposit feeders and suspension feeders in the biotope.
Where the biotope occurs in the shallow subtidal, it is likely to be affected by winter storms. Storms may cause dramatic changes in 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 success of recruitment by newly settled spat or larvae (see Hall, 1994 for review). For example, during winter gales along the North Wales coast large numbers of Abra alba and Mysella bidentata were cast ashore and over winter survival rate was as low as 7% and 50% respectively in the more exposed locations (Rees et al., 1976). Sediment transport and the risk of smothering also occurs.

Habitat structure and complexity

  • Structural complexity is provided by the many tube building species in the biotope. The principal tube builders are the polychaetes Polydora ciliata and Spiophanes bombyx and the amphipods Ampelisca sp. and Haploops tubicola. The tubes built by Polydora ciliata for example are embedded in the sediment and the ends extend a few millimetres above the substratum surface. The mats of agglomerated sediment may be up to 50 cm thick.
  • High densities of tube builders and the presence of tubes favours further sedimentation of fine particles (e.g. Mills (1967) for Ampelisca vadorum and Ampelisca abdita) and may be a factor in stimulating recruitment of species such as Haploops tubicola (Glemarec et al., 1986, cited in Dauvin & Bellan-Santini, 1990).
  • Additional structural complexity is provided by the burrows of infauna although these are generally simple. Most species living within the sediment are limited to the area above the anoxic layer, the depth of which will vary depending on sediment particle size and organic content. However, the presence of burrows of species such as Arenicola marina allows a larger surface area of sediment to become oxygenated, and thus enhances the survival of a considerable variety of small species (Pearson & Rosenberg, 1978). Underlying sediments may also become oxygenated by the activities of amphipods within their tubes (Mills, 1967).

Productivity

Production in IMU.TubeAP is mostly secondary, derived from detritus and organic material. Where, the biotope occurs in shallow subtidal waters, some primary production comes from benthic microalgae (microphytobenthos e.g. diatoms, flagellates and euglenoides) and water column phytoplankton. Beyond 30m depth, there is unlikely to be any in situ primary production. In all cases, the benthos is supported predominantly by pelagic production and by detrital materials emanating from the coastal fringe (Barnes & Hughes, 1992). The amount of planktonic food reaching the benthos is related to:
  • depth of water through which the material must travel;
  • magnitude of pelagic production;
  • proximity of additional sources of detritus;
  • extent of water movement near the sea bed, bringing about the renewal of suspended supplies (Barnes & Hughes, 1992).
Food becomes available to deposit feeders by sedimentation on the substratum surface and by translocation from the water column to the substratum through production of pseudofaeces by suspension feeders.
Productivity in the biotope is expected to be high. The amphipods in particular have a short life span, grow to maturity quickly and have multiple generations per year.
The sediment in the biotope may be nutrient enriched due to proximity to anthropogenic nutrient sources such as sewage outfalls or eutrophicated rivers.

Recruitment processes

  • The spawning period for Polydora ciliata in northern England is from February until June and three or four generations succeed one another during the spawning period (Gudmundsson, 1985). After a week, the larvae emerge and are believed to have a pelagic life from two to six weeks before settling (Fish & Fish, 1996). The larvae settle preferentially on substrates covered with mud (Lagadeuc, 1991).
  • The mating system of amphipods is polygynous and several broods of offspring are produced, each potentially fertilised by a different male. There is no larval stage and embryos are brooded in a marsupium, beneath the thorax. Embryos are released as subjuveniles with incompletely developed eighth thoracopods and certain differences in body proportions and pigmentation. Dispersal is limited to local movements of these subjuveniles and migration of the adults and hence recruitment is limited by the presence of local, unperturbed source populations (Poggiale & Dauvin, 2001). Dispersal of subjuveniles may be enhanced by the brooding females leaving their tubes and swimming to uncolonized areas of substratum before the eggs hatch (Mills, 1967).
  • The tube building polychaetes, e.g. Pygospio elegans, generally disperse via a pelagic larval stage (Fish & Fish, 1996) and therefore recruitment may occur from distant populations. However, dispersal of the infaunal deposit feeders, such as Scoloplos armiger and Arenicola marina, occurs through burrowing of the benthic larvae and adults (Beukema & de Vlas, 1979; Fish & Fish, 1996). Recruitment must therefore occur from local populations or by longer distance dispersal during periods of bedload transport. Recruitment is therefore likely to be predictable if local populations exist but patchy and sporadic otherwise.

Time for community to reach maturity

A community containing Polydora ciliata is likely to reach maturity very rapidly because Polydora ciliata is a short lived species that reaches maturity within a few months and has three or four spawnings during a breeding season. For example, in colonization experiments in Helgoland (Harms & Anger, 1983) Polydora ciliata settled on panels within one month in the spring. The tubes built by Polydora ciliata agglomerate sometimes to form layers of mud up to 20cm thick. However, it may take several years for a Polydora ciliata 'mat' to reach a significant size.
The life cycles of amphipods varies between the different families. Based on the intertidal species, Corophium volutator, the Corophium sp. may produce several broods over the summer breeding season (Fish & Fish, 1996). Haploops tubicola produces 1 or 2 broods per year with a longevity of 2 or 3 years (Dauvin & Bellan-Santini, 1990) and Mills (1967) reported that Ampelisca vadorum and Ampelisca abdita produced only 1 brood per generation but there were 2 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).

Additional information

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Preferences & Distribution

Recorded distribution in Britain and IrelandCommon in lagoons on Shetland and Orkney. No records on mainland coasts of Britain or Ireland.

Habitat preferences

Depth Range
Water clarity preferences
Limiting Nutrients No information found
Salinity
Physiographic
Biological Zone
Substratum
Tidal
Wave
Other preferences

Additional Information

Species composition

Species found especially in this biotope

  • Haploops tubicola

Rare or scarce species associated with this biotope

-

Additional information

Sensitivity reviewHow is sensitivity assessed?

Explanation

The tube building polychaetes Polydora ciliata and Spiophanes bombyx form a key component of the biotope through stabilizing the sediment and encouraging faunal diversity. Loss of these species would probably result in a much more uniform habitat dominated by infaunal deposit feeding polychaetes. The amphipods, either Haploops tubicola or Ampelisca sp. (depending on environment) and Corophium sp. are characteristic of the biotope. Loss of these species would result in loss of the biotope but would not necessarily have cascading effects on the community. However, there are probably important effects on the trophic link between these benthic prey species and the demersal fish which are strong predators.

In the absence of complete Biology and Sensitivity Key Information Reviews for the amphipods that characterize this biotope, the review of the tube dwelling amphipod Jassa falcata has been used for reference.

Species indicative of sensitivity

Community ImportanceSpecies nameCommon Name
Important characterizingAmpelisca spp.An amphipod
Important characterizingCorophium spp.An amphipod
Important characterizingHaploops tubicolaAn amphipod
Key functionalPolydora ciliataA bristleworm
Key functionalSpiophanes bombyxA bristleworm

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High High Moderate Major decline High
Removal of the substratum would remove the entire benthic population and the tubes of the tube building species. A portion of the amphipod population would probably be able to escape the substratum loss through swimming but the majority of other species in the biotope would be lost and there would be a major decline in species richness. Recoverability is recorded as high (see additional information below).
Intermediate High Low No change Low
Smothering with 5 cm of sediment would cover amphipod tubes and prevent suspension feeding. Some mortality of the population is likely. The tubes of polychaetes, including Polydora ciliata, would also be covered and the population would have to build new tubes at the new sediment surface, with some energetic cost. The infaunal burrowing polychaetes would probably be able to relocate to their preferred depth and hence are unlikely to be intolerant. Based on the intolerance of the amphipods, the biotope intolerance is assessed as intermediate but there is unlikely to be a decline in species richness. Recoverability is recorded as high (see additional information below).
Low Very high Very Low No change Low
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 was initiated by the turbidity of the water surrounding the tubes. However, the feeding structures of suspension feeders such as Ampelisca sp. and Haploops tubicola may become clogged by large increases in suspended sediment or feeding may be terminated, compromising growth. Intolerance is therefore assessed as low. Growth would quickly return to normal when suspended sediment returns to original levels so recoverability is recorded as very high.
Tube building polychaetes are not likely to be intolerant 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 is able to tolerate different levels of suspended solids (Read et al., 1982; Read et al., 1983).
Low Very high Moderate Minor decline Very low
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. The benchmark change only lasts for a month and so mortality is unlikely. Intolerance is therefore assessed as low. Growth would quickly return to normal when suspended sediment returns to original levels so recoverability is recorded as very high.
Not relevant Not relevant Not relevant Not relevant Not relevant
The biotope occurs below 5 m depth and therefore desiccation is never likely to be a relevant factor.
Not relevant Not relevant Not relevant Not relevant Not relevant
The biotope occurs below 5 m depth and therefore change in emergence regime is never likely to be a relevant factor.
Not sensitive* Not relevant
The biotope occurs below 5 m depth and therefore change in emergence regime is never likely to be a relevant factor.
Intermediate High Low Decline Low
The biotope occurs in areas of 'weak' tidal streams (Connor et al., 1997a) and is therefore likely to be intolerant of increases in water flow to some degree. An increase in water flow of 2 categories would place the biotope in areas of 'strong' flow. The increase would change the sediment characteristics in which the biotope occurs, primarily by re-suspending and preventing deposition of finer particles (Hiscock, 1983). The underlying sediment in the biotope has a high silt content; a substratum which would not occur in very strong tidal streams. There would be a decrease in tube building material 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. A biotope intolerance of intermediate is therefore recorded and species richness is expected to decline. Recoverability is assessed as high (see additional information below).
Tolerant Not sensitive* No change Low
The biotope occurs in areas of 'weak' tidal streams (Connor et al., 1997a), the characterizing species are adapted to low flow conditions and hence the biotope is unlikely to be intolerant of a further reduction in water flow.
Intermediate High Low Minor decline Low
Murina (1997) categorised 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 global distribution. Amphipods, however, are reported to have low tolerance to temperature changes (Bousfield, 1973) although lethal limits are not given. Biotope intolerance is therefore recorded as intermediate and recoverability as high (see additional information below). Some amphipod species may be lost with a consequent minor decline in species richness. The infauna in the biotope are likely to be insulated from extreme changes of temperature.
Intermediate High Low Minor decline Low
Murina (1997) categorised 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 global distribution. The species survived a drop in temperature from 11.5 to 7.5°C over the course of 15 hours (Gulliksen, 1977) and so it appears the species is tolerant of acute temperature decreases. During the extremely cold winter of 1962/63 when temperatures dropped below freezing point for several weeks, Polydora ciliata was apparently unaffected (Crisp (ed.), 1964). Amphipods, however, are reported to have low tolerance to temperature changes (Bousfield, 1973) although lethal limits are not given. Mills (1967) reported that gonadal growth of Ampelisca vadorum and Ampelisca abdita is retarded by low temperatures, thus delaying maturity, and feeding rate was reduced below 10°C. Biotope intolerance is therefore recorded as intermediate and recoverability as high (see additional information below). Some amphipod species may be lost with a consequent minor decline in species richness. The infauna in the biotope are likely to be insulated from extreme changes of temperature.
Low Very high Very Low No change Low
The biotope occurs in relatively turbid waters and therefore the species in the biotope are likely to be well adapted to turbid conditions. An increase in turbidity may affect primary production in the water column and therefore reduce the availability of diatom food, both for suspension feeders and deposit feeders. In addition, primary production by the microphytobenthos on the sediment surface may be reduced, further decreasing food availability for deposit feeders. However, primary production is probably not a major source of nutrient input into the system and, furthermore, phytoplankton will also immigrate from distant areas so the effect may be decreased. As the benchmark turbidity increase only persists for a year, decreased food availability would probably only affect growth and fecundity of the intolerant species so a biotope intolerance of low is recorded. As soon as light levels return to normal, primary production will increase and hence recoverability is recorded as very high.
Tolerant Not sensitive* No change High
A decrease in turbidity will mean more light is available for photosynthesis by phytoplankton in the water column and microphytobenthos on the sediment surface. This would increase the primary production in the biotope and may mean greater food availability for deposit feeders and suspension feeders. However, primary production is probably not a major source of production in the biotope so the turbidity decrease is not likely to have a significant effect.
High High Moderate Decline Low
The biotope occurs in 'sheltered' and 'very sheltered' areas (Connor et al., 1997a). This suggests that the biotope would be intolerant of wave exposure to some degree. An increase in wave exposure by two categories for one year would be likely to affect the biotope in several ways. Fine sediments would be eroded (Hiscock, 1983) resulting in the likely reduction of the habitat of the infaunal species, a decreased supply of tube building material and a decrease in food availability for deposit feeders. Furthermore, strong wave action is likely to cause damage or withdrawal of delicate feeding and respiration structures of species within the biotope resulting in loss of feeding opportunities and compromised growth. Mills (1967) reported that Ampelisca flats in Barnstable, USA, were damaged noticeably by winter storms. It is likely that high mortality would result and therefore an intolerance of high is recorded and species richness is expected to decline. Recoverability is recorded as high (see additional information below).
Tolerant Not sensitive* No change Low
The biotope occurs in 'sheltered' and 'very sheltered' areas (Connor et al., 1997a). A decrease in wave exposure by 2 categories for a year would place a portion of the biotope in 'ultra sheltered' areas. The characterizing species are adapted to low flow conditions and are unlikely to be intolerant of this change. The consequent increased risk of smothering is detailed in the relevant section.
Tolerant Not relevant Not relevant No change High
There is no evidence to suggest that any of the species which characterize the biotope are sensitive to noise or vibration at the level of the benchmark.
Low Very high Very Low No change Moderate
Polydora ciliata responds to visual disturbance by withdrawing its palps into its burrow, believed to be a defence against predation. Since the withdrawal of the palps interrupts feeding and possibly respiration the species also shows habituation of the response (Kinne, 1970). Growth may be compromised by the interruption of feeding and so intolerance is assessed as low. Growth should quickly return to normal when the disturbance is over so recoverability is recorded as very high.
Intermediate High Low No change Low
Many species in the biotope are vulnerable to physical abrasion. The tubes of the polychaetes and amphipods are bound only with mucous and are therefore likely to damaged or removed by the benchmark level of abrasion (a scallop dredge). The soft bodied polychaetes are most likely to suffer mortality, while the more robust amphipods are likely to be more resistant, and mobile enough to avoid impact. The infaunal annelids are predominantly soft bodied, live within a few centimetres of the sediment surface and may expose feeding or respiration structures where they could easily be damaged by a physical disturbance such as a dredge. Biotope intolerance is therefore recorded as intermediate. Recoverability is recorded as high (see additional information below).
Low Very high Very Low No change Moderate
Polydora ciliata is capable of tube building throughout its life and so is able to re-establish itself following displacement. In experimental removal of Polydora ciliata, individuals of all ages which were removed from their tubes all built new tubes (Daro & Polk, 1973). Similarly, amphipods are mobile animals which would be able to build new tubes or burrows following displacement. Time and energy would have to diverted to the tube building process at the expense of growth and reproduction so intolerance is assessed as low. Following displacement, the growth would quickly return to normal so recoverability is assessed as very high.

Chemical Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
High High Moderate Decline High
The tube building polychaetes appear to be not sensitive to synthetic chemicals. For example, spionid polychaetes were found by McLusky (1982) to be relatively tolerant of distilling and petrochemical industrial waste in Scotland and Polydora ciliata was abundant at polluted sites close to acidified, halogenated effluent discharge from a bromide-extraction plant in Amlwch, Anglesey (Hoare & Hiscock, 1974). However, gammaridean amphipods have been reported to be intolerant of tri-butyl tin (TBT), a previously common component of antifouling paints, with 10 day LC50 values of 1-48 ng/l (Meador et al., 1993). Biotope intolerance is therefore assessed as high. Recoverability is recorded as high (see additional information below) but is likely to depend on the length of time that synthetic chemicals persist in the sediment. Beaumont et al. (1989) concluded that TBT had a detrimental effect on the larval and/or juvenile stages of infaunal polychaetes and Arenicola marina was found to be intolerant of ivermectin through the ingestion of contaminated sediment (Thain et al., 1998; cited in Collier & Pinn, 1998). Beaumont et al. (1989) also concluded that bivalves are particularly intolerant of TBT. It is expected therefore that there would be a decline in species richness following exposure to synthetic chemicals.
Heavy metal contamination
Intermediate Moderate Moderate Decline Moderate
Experimental studies with various species suggests that polychaete worms are quite tolerant of heavy metals (Bryan, 1984). Crustaceans are generally regarded to be intolerant of cadmium (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. Biotope intolerance is therefore assessed as intermediate and, since heavy metals are likely to persist in sediments, recoverability as moderate (see additional information below). Bivalves are also intolerant of heavy metal contamination (e.g. Eisler, 1977; Kaschl & Carballeira, 1999) so species richness in the biotope would be expected to decline.
Hydrocarbon contamination
High Low High Decline High
Amphipods in general, and ampeliscid amphipods in particular seem particularly intolerant of contamination with oil. Dauvin (1998) reported the effects of the Amoco Cadiz oil spill on the fine sand Abra alba community in the Bay of Morlaix. Reductions in abundance, biomass and production of the community were very evident through the disappearance of the dominant populations of the amphipods Ampelisca sp. The spill occurred in 1978 and after 2 weeks, the level of hydrocarbons in subtidal sediments reached 200 ppm (Dauvin, 1984; cited in Poggiale & Dauvin, 2001). This caused the disappearance of the Ampelisca populations, leaving behind a single species, Ampelisca sarsi, in very low densities. The sediment rapidly depolluted and in 1981 benthic recruitment occurred in normal conditions (Dauvin, 1998). However, the recovery of the Ampelisca populations took up to 15 years. Intolerance is therefore recorded as high with a decline in species richness. In view of the recovery of the amphipod populations, recoverability is assessed as low.
Radionuclide contamination
No information Not relevant No information Insufficient
information
Not relevant
No evidence was found concerning the intolerance of the characterizing species to radionuclide contamination.
Changes in nutrient levels
Tolerant Not relevant Not relevant No change High
Polydora ciliata is often found in environments subject to high levels of nutrients (Sordino et al., 1989). For example, the species was abundant in areas of the Firth of Forth exposed to high levels of sewage pollution (Smyth, 1968) and, in an organically polluted fjord receiving effluent discharge from Oslo, Polydora ciliata settled in large numbers within the first month (Green, 1983, Pardal et al., 1993). Similarly, amphipods appear to be tolerant of, and indeed prefer, high nutrient levels. Haploops tubicola muds, for example, are indicative of organically enriched sediments (Le Bris & Glemarec, 1995). The biotope is therefore assessed as not sensitive.
Not relevant Not relevant Not relevant Not relevant Not relevant
The biotope occurs in full salinity conditions and is therefore not likely to be subject to increases in salinity. No information was found concerning the reaction of the characterizing species to hypersaline conditions.
Low Very high Moderate Minor decline Very low
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). Of the amphipod species, Corophium has a very high tolerance of low salinity (Fish & Fish, 1996) but this might only be true of the species with estuarine distributions. It is likely that the marine species would be intolerant of salinity decreases in some way and so biotope intolerance is assessed as low. Some species in the biotope are highly intolerant of reduced salinity. For example, Arenicola marina is unable to tolerate salinities below 24 psu and is excluded from areas influenced by freshwater runoff or input (e.g. the head end of estuaries) (Hayward, 1994). There is therefore likely to be a minor decline in species richness in the biotope.
High High Moderate Decline Moderate
Amphipods appear not to be tolerant of reduced oxygenation. Ampelisca agassizi, for example, is reported to be intolerant of hypoxia (see review by 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). There is therefore likely to be some mortality of amphipods after a week at the benchmark level of hypoxia. Polydora ciliata is apparently tolerant of hypoxia as the species is repeatedly found at localities with oxygen deficiency (Pearson & Rosenberg, 1978). For example, in polluted harbours in Los Angeles and Long Beach, 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). In view of the intolerance of amphipods, biotope intolerance is assessed as high and there would be a decline in species richness. Recoverability is recorded as high (see additional information below).

Biological Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
No information Not relevant No information Insufficient
information
Not relevant
No information was found concerning the infection of the characterizing species by microbial pathogens.
No information Not relevant No information Insufficient
information
Not relevant
There is no information to suggest that the biotope is threatened by invasion of alien species.
Not relevant Not relevant Not relevant Not relevant Not relevant
It is extremely unlikely that any of the species indicative of sensitivity would be targeted for extraction and we have no evidence for the indirect effects of extraction of other species on this biotope.
Not relevant Not relevant Not relevant Not relevant Not relevant

Additional information

Recoverability
Amphipods have a short life span, mature quickly and may have multiple generations per year (Mills, 1967; Dauvin & Bellan-Santini, 1990) suggesting that they would have strong powers of recoverability. However, fecundity is generally low, there is no larval stage and the embryos are brooded in a marsupium, beneath the thorax. Dispersal is limited to local movements of the subjuveniles and migration of the adults and hence recruitment is limited by the presence of local, unperturbed source populations (Dauvin, 1998). Poggiale & Dauvin (2001) reported that recovery of an Ampelisca population took up to 15 years, but this was following an oil spill, to which amphipods are particularly intolerant, and it is likely that this is an exceptional situation. It is expected that, in situations where there is no residual population, amphipods would normally recover within 5 years and so recoverability is assessed as high.
The tube building polychaetes, including Polydora ciliata, are moderately fecund, the planktonic larvae are capable of dispersal over long distances and the reproductive period is of several months duration. In colonization experiments in Helgoland, Polydora ciliata settled on panels within one month in the spring (Harms & Anger, 1983). Recovery and establishment of a mature community is likely to occur within 5 years and so recoverability is assessed as high.
Based on the recoverability of the characterizing species, biotope intolerance is assessed as high.

Importance review

Policy/Legislation

Habitats of Principal ImportanceSubtidal sands and gravels
Habitats of Conservation ImportanceSubtidal sands and gravels
Habitats Directive Annex 1Sandbanks which are slightly covered by sea water all the time
UK Biodiversity Action Plan PrioritySubtidal sands and gravels

Exploitation

No species found within this biotope are known to be exploited.

Additional information

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Citation

This review can be cited as:

Rayment, W.J. 2002. Semi-permanent tube-building amphipods and polychaetes in sublittoral sand. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from: http://www.marlin.ac.uk/habitat/detail/136

Last Updated: 20/03/2002