Aphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud

Researched byDr Keith Hiscock Refereed byThis information is not refereed.
EUNIS CodeA5.322 EUNIS NameAphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud

Summary

UK and Ireland classification

EUNIS 2008A5.322Aphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud
EUNIS 2006A5.322Aphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud
JNCC 2004SS.SMu.SMuVS.AphTubiAphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud
1997 BiotopeSS.IMU.EstMu.AphTubAphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud

Description

Variable salinity cohesive muddy sediment dominated by the polychaete Aphelochaeta marioni and the oligochaetes Tubificoides spp. The polychaetes Polydora ciliata, Cossura longocirrata and Melinna palmata may also occur in high numbers. The cirratulid polychaete Caulleriella zetlandica may also occur (there is still inconsistency in the identification of the cirratulid group, compounded by fragmentation during sample processing). This biotope is very common in stable muddy environments and may extend from reduced salinity to fully marine conditions. The biotope may be separated from similar biotopes such as IMU.NhomTub by the abundance of Aphelochaeta marioni, terebellids and an indication of the stability of the sediment. In areas of mixed sediment Aphelochaeta marioni may also occur in high numbers. In this case it may be difficult to separate IMU.AphTub from IMX.PolMtru requiring classification on sediment characteristics and associated species such as the bivalve Mya truncata in addition to the abundance of Aphelochaeta marioni. It may be separated from IMX.CreAph by the relative abundances of the slipper limpet Crepidula fornicata in addition to Aphelochaeta marioni. This biotope may also be found in conjunction with IMS.MacAbr. (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

Recorded from estuarine habitats in England and in south Wales.

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 or burrow-living polychaetes and by oligochaetes, with errant polychaetes foraging in the surrounding and underlying sediment.
  • Mobile, carnivorous polychaetes, including Nephtys hombergi, Anaitides spp, Eteone longa, and Pholoe spp., predate the smaller annelids and crustaceans.
  • The dominant tube-builders are the deposit feeding polychaetes Polydora ciliata and Lanice conchilega. 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 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).
  • In some examples of the biotope, the tube-building, suspension feeding amphipod Ampelisca sp. and the burrowing Corophium volutator are present
  • 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).
  • Some examples of the biotope contain a number of infaunal bivalve species, including Abra alba, Abra nitida and Mysella bidentata, which probably both deposit feed and suspension feed depending on local environmental conditions.
  • Foraging species such as Carcinus maenas and Crangon crangon may feed selectively and influence the composition of the biotope.

Seasonal and longer term change

The biotope is present throughout the year with the possibility of some seasonal variation in numbers of each species. Hall & Frid (1998) found that colonization by many of the polychaetes associated with this biotope did not vary significantly with season although recruitment of Tubificoides benedii and Ophyrotrocha hartmanni did vary significantly with season.

Habitat structure and complexity

  • Structural complexity is provided by the many tube building species in the biotope. 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.
  • 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. Underlying sediments may also become oxygenated by the activities of amphipods within their tubes (Mills, 1967), burrowing bivalves and polychaetes.

Productivity

Production in IMU.Aph.Tub 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. 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, and the
  • 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. Many of the characterizing species are likely to 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

Limited information has been found on species in the biotope and only characterizing species have been specifically researched.
  • The lifecycle of Aphelochaeta marionivaries according to environmental conditions. In Stonehouse Pool, Plymouth, Aphelochaeta marioni (studied as Tharyx marioni) spawned in October and November (Gibbs, 1971) whereas in the Wadden Sea, Netherlands, spawning occurred from May to July (Farke, 1979). The embryos developed lecithotrophically and hatched in about 10 days (Farke, 1979). Under stable conditions, adult and juvenile Aphelochaeta marioni will disperse by burrowing (Farke, 1979).
  • 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).
  • Nephtys hombergi exhibits variable spawning success with failures in some years (Olive et al., 1997).
  • The mating system of amphipods is polygynous and several broods of offspring are produced, each potentially fertilized by a different male. There is no larval stage and embryos are brooded in a marsupium, beneath the thorax. Embryos are released as sub-juveniles with incompletely developed eighth thoracopods and certain differences in body proportions and pigmentation. 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 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 (Boström & Bonsdorff, 2000). However, dispersal of some of the infaunal deposit feeders, such as Scoloplos armiger, 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

The community is dominated by fast growing opportunistic species and the community most likely reaches maturity within one year of space becoming available. In an experimental study investigating recovery of a range of species characteristically found in this biotope after copper contamination, Hall & Frid (1995) found that recovery took up to a year. However, Hall & Frid (1998) found that colonization by many of the polychaetes associated with this biotope did not vary significantly with season although recruitment of Tubificoides benedii and Ophyrotrocha hartmanni did vary significantly with season.

Additional information

None

Preferences & Distribution

Recorded distribution in Britain and IrelandRecorded from estuarine habitats in England and in south Wales.

Habitat preferences

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

Additional Information

None

Species composition

Species found especially in this biotope

    Rare or scarce species associated with this biotope

    -

    Additional information

    Nephtys hombergi and Tubificoides spp. have also been researched as a part of the sensitivity assessment although separate reviews have not yet (April 2002) been prepared. Four hundred and seventy two species have been recorded from this biotope by the Marine Nature Conservation Review (JNCC, 1999) although many in small numbers or in few examples.

    Sensitivity reviewHow is sensitivity assessed?

    Explanation

    The species listed are ones that commonly occur in the biotope and represent a range of functional types. Polydora ciliata gives structure to the biotope by stabilizing sediments and may also be a dominant species excluding others. Aphelochaeta marioni is characteristic of the biotope. Hediste diversicolor occurs in the biotope but has been included in the species representative of sensitivity to represent similar polychaetes such as Nephtys hombergi.

    Species indicative of sensitivity

    Community ImportanceSpecies nameCommon Name
    Important characterizingAphelochaeta marioniA bristleworm
    Important otherHediste diversicolorRagworm
    Important otherHydrobia ulvaeLaver spire shell
    Important otherLanice conchilegaSand mason worm
    Key structuralPolydora ciliataA bristleworm

    Physical Pressures

     IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
    High High Moderate Major decline High
    Removal of the substratum would remove the entire benthic population. Significant recolonization by many species in the biotope might occur within a few months but the biotope would be unlikely to be recognised until after six months. Recoverability is therefore recorded as high (see additional information below).
    Intermediate Very high Low No change High
    The characterizing species are all mobile and capable of burrowing through 5 cm of smothering sediment. Some mortality of the population may, however occur. Tube building polychaetes, including Polydora ciliata, would be covered and the population would have to build new tubes at the new sediment surface, with some energetic cost. Hydrobia ulvae may not be able to reach the sediment surface. The infaunal burrowing polychaetes would probably be able to relocate to their preferred depth and hence are unlikely to be sensitive. Based on the likelihood that some individuals of some species would perish, the biotope intolerance is assessed as intermediate but there is unlikely to be a decline in species richness. Recoverability is recorded as very high (see additional information below).
    Tolerant* Not relevant Not sensitive* No change High
    The biotope occurs in estuarine waters that are subject to occasional very high suspended sediment loads. Most of the species in this biotope are deposit feeders and may benefit from increased settlement of detritus from increased siltation. Tube building polychaetes are likely to tolerate high suspended sediment 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). The biotope may benefit from an increase in suspended sediment.
    Low High Moderate Major decline 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 from the lowest shore downwards and may be subject to desiccation. However, all of the characterizing species are burrowing and other frequently occurring species that are surface dwellers may be able to migrate (for instance: Crangon crangon, Hydrobia ulvae, Carcinus maenas). Therefore, desiccation, where the biotope occurs on the lower shore upward extent of its range, is considered not relevant.
    Low Very high Very Low No change Low
    The biotope occurs from the lowest shore downwards and may be subject to significant desiccation if emergence increased. Also, during heavy rain, low salinity is a consideration. However, all of the characterizing species are burrowing and other frequently occurring species that are surface dwellers may be able to migrate (for instance: Crangon crangon, Hydrobia ulvae, Carcinus maenas). The biotope occurs in situations subject to variable salinity and species are protected within the sediment and fairly stable interstitial water salinity and not expected to be intolerant of occasional downpours. Therefore, a minority of the community would be expected to be affected by increased emergence and no great alteration to the abundance of dominant or characterizing species so that an intolerance of low is suggested but with low confidence. Overall, the biotope would not be changed and so a recoverability of very high is suggested (see additional information below).
    Tolerant* Not sensitive No change Moderate
    The biotope is predominantly subtidal and a decrease in emergence would be unlikely to have any adverse effect and would increase the habitat available for development of the biotope.
    High High Moderate Decline Low
    The biotope occurs in areas of 'weak' to 'moderately strong' tidal streams (Connor et al., 1997b) and is therefore likely to be intolerant of increases in water flow to some degree. An increase in water flow of 2 categories could place the biotope in areas of 'very strong' flow. Although muddy sediments are cohesive and may resist winnowing by strong currents, the turbulence involved in tidal flows of 3 knots and more will most likely alter the substratum. The increase would change the sediment characteristics in which the biotope occurs, primarily by re-suspending and preventing deposition of finer particles (Hiscock, 1983). 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. Overall, the biotope is likely to change to one that is characteristic of coarser sediments. A biotope intolerance of high is therefore recorded and species richness is expected to decline. Recoverability is assessed as high (see additional information below) especially as silt, from typically high turbidity estuarine conditions, is likely to redeposit rapidly.
    Tolerant Not sensitive* No change Moderate
    The biotope occurs in areas of 'weak' tidal streams (Connor et al., 1997b), 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. (The possibility of water becoming stagnant and, because wave action is typically very low in this biotope, de-oxygenated is considered later in 'Changes in oxygenation'.)
    Low Very high Very Low No change Low
    Bamber & Spencer (1984) observed that Tubificoides and Caulleriella species, common species in the biotope, were dominant in the area affected by thermal discharge in the River Medway estuary. Murina (1997) categorised Polydora ciliata as a eurythermal species because of its ability to spawn in temperatures ranging from 10.6-19.9° C. Increased temperature may have indirect effects. For instance, higher temperatures have been implicated in the proliferation of trematode parasites which have caused mass mortalities in the snail Hydrobia ulvae (Jensen & Mouritsen, 1992). No other information has been found on tolerance of component species to increased temperature although it would be expected that the infauna in the biotope will be insulated from extreme changes of temperature. Nevertheless, an increase in temperature may indirectly affect some species as microbial activity within the sediments will be stimulated increasing oxygen consumption and promoting hypoxia (see 'Change in oxygenation' below). An intolerance of low is suggested but with a low confidence. Recoverability is likely to be rapid.
    Low Immediate Not relevant No change Low
    Very little information has been found describing the tolerance of component species in the biotope to low temperatures. Beukema et al. (1988) observed that Nephtys hombergi showed a lower survival in the (colder) north-east part of the Wadden Sea compared to the south-west. Polydora ciliata 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, 1964). Observations in Crisp (1964) described mortality of Lanice conchilega between the tidemarks but not at lower levels. However, species dwelling in the sediments (at the upper intertidal limits of this biotope) are likely to be protected from the direct effects of temperature change at the surface. For instance, Hediste diversicolor burrows deeper in very cold and frosty weather (Linke, 1939). Overall, although mortality seems unlikely, especially as the biotope is mainly subtidal, some reduction in feeding and loss of condition may occur and an intolerance of low has been reported. Recovery would be likely to be immediate.
    Low Very high Very Low No change Moderate
    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 Moderate
    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 Major decline Moderate
    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. 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 Moderate
    The biotope occurs in 'sheltered' and 'very sheltered' areas (Connor et al., 1997b). For a subtidal biotope, there is therefore likely to be very little oscillatory water movement and the predominant water movement will be tidal flow. 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 likely to tolerate this change.
    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
    Some of the species in the biotope may be intolerant of shading but would not 'see' predators. Farke (1979) noted their intolerance of Aphelochaeta marionito disturbance by light in a microsystem in the laboratory. Polydora ciliata responds to shading by withdrawing its palps into its burrow, believed to be a defence against predation (Kinne, 1970). Although not strictly "visual presence", the withdrawal of feeding structures means that 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 Very high Low Decline Low
    Many species in the biotope are vulnerable to physical abrasion. The tubes of the polychaetes are bound only with mucous and are therefore likely to damaged by a passing scallop dredge. 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. Biotope intolerance is therefore recorded as intermediate. Recoverability is recorded as very high as damage at the benchmark level will be restricted in extent (see additional information below). For large scale physical disturbance, sensitivity will be more similar to 'substratum removal' above.
    High Moderate Moderate Decline Low
    The species in the biotope are either mobile and capable of re-burrowing or, mainly, capable of re-building tubes. However, following displacement of key or characterizing species, the biotope would have to be structurally re-established - there may be a succession of species before IMU.AphTub is recognised. Intolerance is identified as high and recoverability moderate but with low confidence.

    Chemical Pressures

     IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
    Intermediate High Low Decline Moderate
    Some species in the biotope are known to be adversely affected by synthetic chemicals. For instance, Scoloplos armiger (frequently found in the biotope) exhibited 'moderate' intolerance to tri-butyl tin antifoulants (Bryan & Gibbs, 1991). Collier & Pinn (1998) investigated the effect on the benthos of Ivermectin, a feed additive treatment for infestations of sea-lice on farmed salmonids. The polychaete Hediste diversicolor(frequently found in the biotope) was particularly susceptible, exhibiting 100% mortality within 14 days when exposed to 8 mg/m² of Ivermectin in a microcosm. On the other hand, Beaumont et al. (1989) investigating the effects of tri-butyl tin (TBT) on benthic organisms found that at concentrations of 1-3 µg/l there was no significant effect on the abundance of Hediste diversicolor or Cirratulus cirratus (an infrequent component of the biotope) after 9 weeks in a microcosm. However, no juvenile polychaetes were retrieved from the substratum and hence there is some evidence that TBT had an effect on the larval and/or juvenile stages of these polychaetes. 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, oligochaetes (principally Tubificoides benedeni) and Hydrobia ulvae were found by McLusky (1982) to be amongst the most tolerant species in the vicinity of a of a petrochemical industrial waste in the Firth of Forth, Scotland. The biotope occurs in polluted conditions and overall, an intolerance of intermediate is suggested reflecting the likelihood that some chemicals might adversely affect some species reducing abundance and viability but the biotope would persist. For recoverability, see additional information. Recovery would require synthetic chemicals to have depurated from the sediment.
    Heavy metal contamination
    Intermediate High Low Decline Low
    The majority of species in this biotope are polychaetes and evidence suggests that they are "fairy resistant" to the effects of heavy metals (Bryan, 1984). However, Hall & Frid (1995) found that the four dominant taxa in their study (species typically found in this biotope including Tubificoides spp. and Capitella capitata) were reduced in abundance in copper-contaminated sediments and that recovery took up to one year after the source of contamination ceased. Some other species (for instance Carcinus maenas) , may adapt to high metal concentrations (Bryan, 1984). Polydora ciliata, one of the species that occurs frequently in the biotope, occurs 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). However, Hediste diversicolor has been found successfully living in estuarine sediments contaminated with copper ranging from 20 µm Cu/g in low copper areas to >4000 µm Cu/g where mining pollution is encountered e.g. Restronguet Creek in the Fal Estuary, Cornwall (Bryan & Hummerstone, 1971). Taking account of the low salinity conditions that affect this biotope (in general, for estuarine animals, heavy metal toxicity increases as salinity decreases and temperature increases: McLusky et al., 1986), it seems possible that some species at least in the biotope might be adversely affected by high contamination by heavy metals. The assessment of intermediate intolerance is 'precautionary' and the specific levels at a location would need to be matched to experimental or field studies to assign a more accurate rank. For recoverability, see additional information below. Recovery of species in the biotope would be influenced by the length of time it would take for the habitat to return to a suitable state (e.g. factors such as the decline of bioavailable metals within the marine environment), recolonization by adult and juvenile specimens from adjacent habitats, and the establishment of a breeding population.
    Hydrocarbon contamination
    Intermediate High Low Decline Moderate
    The biotope is predominantly subtidal and component species are protected from the direct effects of oil spills by their depth but are likely to be exposed to the water soluble fraction of oils and hydrocarbons, or hydrocarbons adsorbed onto particulates. Some of the polychaetes in this biotope proliferate after oil spills: for instance Capitella capitata (Suchanek, 1993) and Aphelochaeta marioni (Dauvin, 1982, 2000). Cirratulids seem mostly immune probably because their feeding tentacles are protected by mucus (Suchanek, 1993). Nevertheless it might be expected that some of the species in the biotope may be affected and the increase in abundance of some species suggests reduced competition with others. However, because some species in the biotope may increase in abundance following a spill, and because of the subtidal character of the biotope, it is expected that adverse effects from hydrocarbons may reduce abundance and viability of some species but the biotope would persist. An intolerance of intermediate is therefore suggested but with a high recoverability (see additional information below).
    Radionuclide contamination
    No information Not relevant No information Insufficient
    information
    Not relevant
    No information has been found.
    Changes in nutrient levels
    Low Immediate Not sensitive No change Very low
    It would be expected that some increase in nutrients would favour the expansion of food resources for deposit feeders. Increased nutrients often derive from sewage inputs and presence of species such as Aphelochaeta marioni in such situations (for instance Broom et al., 1991) may reflect tolerance to high nutrients or to deoxygenated conditions or both. Overall, the benefits (higher food resources) and disbenefits (possible hypoxia) make it difficult to determine intolerance but, considering the often eutrophic situations the biotope occurs in, an intolerance of low is suggested but with very low confidence.
    Not relevant Not relevant Not relevant Not relevant Not relevant
    The biotope occurs in reduced to full salinity and so increase in salinity is considered not relevant.
    High High Intermediate Major decline Low
    The biotope occurs in reduced salinity. One of the characterizing species, Aphelochaeta marioni, has been recorded from brackish inland waters in the Southern Netherlands with a salinity of 16 psu, but not in areas permanently exposed to lower salinities (Wolff, 1973). However, it also penetrates into areas exposed to salinities as low as 4 psu for short periods at low tide when fresh water discharge from rivers is high (Farke, 1979). The distribution of Aphelochaeta marioni, therefore, suggests that it is very tolerant of low salinity conditions and would be tolerant of reduced salinity especially for short periods. However, a long term reduction from reduced to low salinity may affect some of the species in the biotope with possible losses and reduced viability. The biotope would probably change to one more tolerant of very low salinity conditions. An intolerance of high is therefore suggested but recovery would be rapid on return to previous conditions (see additional information below).
    Intermediate High Low Minor decline Moderate
    Some of the species frequently found in the biotope (Malacoceros fuliginosus, Nephtys hombergi, Heteromastus filiformis) are noted by Diaz & Rosenberg (1995) as resistant to severe hypoxia or (Capitella capitata, Hediste diversicolor) to moderate hypoxia. Tubificoides benedii has a high capacity to tolerate anoxic conditions (see Giere et al., 1999). Broom et al. (1991) found communities with species characteristic of this biotope in the Severn Estuary where the oxygenated layer was very thin probably as a result of sewage input and suggested that Aphelochaeta marioni was characteristic of faunal assemblages in the Severn Estuary with very poorly oxygenated mud. The successful survival of Hediste diversicolor under prolonged hypoxia was confirmed by the resistance experiments of Vismann (1990), which resulted in a mortality of only 15% during a 22 day exposure of Hediste diversicolor at 10% oxygen (ca. 2.8 mg O2 per litre). Whilst the biotope might thrive in conditions of hypoxia, some species might suffer, reducing species richness. Following a hypoxia event in summer 1994 in the southern Baltic, species (some of which occur in the biotope) took at least two years to recolonize but by summer 1996 had returned to pre-event community structure (Powilleit & Kube, 1999). Since species richness may be reduced by reduction in oxygen, an intolerance of intermediate is suggested reflecting the likelihood that the biotope will not be lost.

    Biological Pressures

     IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
    Low High Low Minor decline Moderate
    No information was found concerning the infection of most of the characterizing species by microbial pathogens. However, there are records of mass mortalities of Hydrobia ulvae caused by high temperatures triggering mass development of larval digenean trematodes within the snails (Jensen & Mouritsen, 1992). The effect on the biotope is likely to be low and recovery high.
    High High Moderate Minor decline Moderate
    Invasion by the slipper limpet Crepidula fornicatamay switch the biotope to IMU.CreAph suggesting high intolerance as the original biotope would be lost. Species richness might decline as Crepidula may dominate the seabed. On the other hand, low densities of Crepidula might have no effect on species richness and add one species (Crepidula) to the community.
    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
    The biotope typically consists of fast growing opportunistic species so that recoverability is expected to be very high or high. However, recovery to full species richness may take longer than one year. The following information has informed the recoverability assessment. Ferns et al. (2000) found that, following significant depletion of Nephtys hombergi by cockle dredging recovery took more than 50 days (but not more than 100 days). Hall & Frid (1998) found that colonization by many of the polychaetes associated with this biotope did not vary significantly with season although recruitment of Tubificoides benedii and Ophyrotrocha hartmanni did vary significantly with season. Also, there may be spawning failure in some years, for instance in Nephtys hombergi (Olive et al. 1997). Following a hypoxia event in summer 1994 in the southern Baltic, species (some of which occur in the biotope) took at least two years to recolonize but by summer 1996 had returned to pre-event community structure (Powilleit & Kube, 1999).

    Importance review

    Policy/Legislation

    Habitats Directive Annex 1Estuaries

    Exploitation

    None known.

    Additional information

    No additional information.

    Bibliography

    1. Bamber, R.N. & Spencer, J.F. 1984. The benthos of a coastal power station thermal discharge canal. Journal of the Marine Biological Association of the United Kingdom, 64, 603-623.
    2. Barnes, R.S.K. & Hughes, R.N., 1992. An introduction to marine ecology. Oxford: Blackwell Scientific Publications.
    3. 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.
    4. 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.
    5. Beukema, J.J., Doerjes, J. & Essink, K., 1988. Latitudinal differences in survival during a severe winter in macrozoobenthic species sensitive to low temperatures. Senckenbergiana maritima, 20, 19-30.
    6. Boström, C. & Bonsdorff, E., 2000. Zoobenthic community establishment and habitat complexity - the importance of seagrass shoot density, morphology and physical disturbance for faunal recruitment. Marine Ecology Progress Series, 205, 123-138.
    7. Broom, M.J., Davies, J., Hutchings, B. & Halcrow, W., 1991. Environmental assessment of the effects of polluting discharges: stage 1: developing a post-facto baseline. Estuarine, Coastal and Shelf Science, 33, 71-87.
    8. Bryan, G.W. & Gibbs, P.E., 1991. Impact of low concentrations of tributyltin (TBT) on marine organisms: a review. In: Metal ecotoxicology: concepts and applications (ed. M.C. Newman & A.W. McIntosh), pp. 323-361. Boston: Lewis Publishers Inc.
    9. Bryan, G.W. & Hummerstone, L.G., 1971. Adaptation of the polychaete Nereis diversicolor to estuarine sediments containing high concentrations of heavy metals. I. General observations and adaption to copper. Journal of the Marine Biological Association of the United Kingdom, 51, 845-863.
    10. 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.
    11. 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.
    12. Connor, D.W., Brazier, D.P., Hill, T.O., & Northen, K.O., 1997b. Marine biotope classification for Britain and Ireland. Vol. 1. Littoral biotopes. Joint Nature Conservation Committee, Peterborough, JNCC Report no. 229, Version 97.06., Joint Nature Conservation Committee, Peterborough, JNCC Report No. 230, Version 97.06.
    13. 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.
    14. Daro, M.H. & Polk, P., 1973. The autecology of Polydora ciliata along the Belgian coast. Netherlands Journal of Sea Research, 6, 130-140.
    15. Dauvin, J.C., 1982. Impact of Amoco Cadiz oil spill on the muddy fine sand Abra alba - Melinna palmata community from the Bay of Morlaix. Estuarine and Coastal Shelf Science, 14, 517-531.
    16. Dauvin, J.C., 2000. The muddy fine sand Abra alba - Melinna palmata community of the Bay of Morlaix twenty years after the Amoco Cadiz oil spill. Marine Pollution Bulletin, 40, 528-536.
    17. 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.
    18. 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.
    19. 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.
    20. Farke, H., 1979. Population dynamics, reproduction and early development of Tharyx marioni (Polychaeta, Cirratulidae) on tidal flats of the German Bight. Veroffentlichungen des Instituts fur Meeresforschung in Bremerhaven, 18, 69-99.
    21. 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.
    22. Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.

    23. Gibbs, P.E., 1971. Reproductive cycles in four polychaete species belonging to the family Cirratulidae. Journal of the Marine Biological Association of the United Kingdom, 51, 745-769.
    24. Giere, O., Preusse, J. & Dubilier, N. 1999. Tubificoides benedii (Tubificidae, Oligochaeta) - a pioneer in hypoxic and sulfide environments. An overview of adaptive pathways. Hydrobiologia, 406, 235-241.
    25. 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.
    26. Gulliksen, B., 1977. Studies from the U.W.L. "Helgoland" on the macrobenthic fauna of rocks and boulders in Lübeck Bay (western Baltic Sea). Helgoländer wissenschaftliche Meeresunters, 30, 519-526.
    27. Hall, J.A. & Frid, C.L.J. 1998. Colonisation patterns of adult macrobenthos in a polluted North Sea Estuary. Aquatic Ecology, 31, 333-340.
    28. Hall, J.A. & Frid, C.L.J., 1995. Response of estuarine benthic macrofauna in copper-contaminated sediments to remediation of sediment quality. Marine Pollution Bulletin, 30, 694-700.
    29. 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.
    30. 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.

    31. 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.
    32. 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,
    33. 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
    34. 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.
    35. Linke, O., 1939. Die Biota des Jadebusenwatts. Helgolander Wissenschaftliche Meeresuntersuchungen, 1, 201-348.
    36. 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.
    37. 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.
    38. 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.
    39. Murina, V., 1997. Pelagic larvae of Black Sea Polychaeta. Bulletin of Marine Science, 60, 427-432.
    40. 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.
    41. Olive, P.J.W. & Cadman, P.S., 1990. Mass mortalities of the lugworm on the South Wales coast: a consequence of algal bloom? Marine Pollution Bulletin, 21, 542-545.
    42. Olive, P.J.W. & Garwood, P.R., 1981. Gametogenic cycle and population structures of Nereis (Hediste) diversicolor and Nereis (Nereis) pelagica from North-East England. Journal of the Marine Biological Association of the United Kingdom, 61, 193-213.
    43. Olive, P.J.W., Porter, J.S., Sandeman, N.J., Wright, N.H. & Bentley, M.G. 1997. Variable spawning success of Nephtys hombergi (Annelida: Polychaeta) in response to environmental variation. A life history homeostasis? Journal of Experimental Marine Biology and Ecology, 215, 247-268.
    44. 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.
    45. Powilleit, M. & Kube, J. 1999. Effects of severe oxygen depletion on macrobenthos of the Pomeranian Bay (southern Baltic Sea): a case study in a shallow, sublittoral habitat characterised by low species richness. Journal of Sea Research, 42, 221-234.
    46. 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.
    47. 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.
    48. Suchanek, T.H., 1993. Oil impacts on marine invertebrate populations and communities. American Zoologist, 33, 510-523.
    49. Vismann, B., 1990. Sulphide detoxification and tolerance in Nereis (Hediste) diversicolor and Nereis (Neanthes) virens (Annelida: Polychaeta). Marine Ecology Progress Series, 59, 229-238.
    50. Wolff, W.J., 1973. The estuary as a habitat. An analysis of the data in the soft-bottom macrofauna of the estuarine area of the rivers Rhine, Meuse, and Scheldt. Zoologische Verhandelingen, 126, 1-242.

    Citation

    This review can be cited as:

    Hiscock, K. 2002. Aphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral 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. Available from: http://www.marlin.ac.uk/habitat/detail/201

    Last Updated: 29/05/2002