Aphelochaeta marioni and Tubificoides spp. in variable salinity infralittoral mud

Researched byEliane De-Bastos & Dr Keith Hiscock Refereed byAdmin
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 (sometimes with some coarser material) dominated by the polychaete Aphelochaeta marioni (or other Aphelochaeta species e.g. Aphelochaeta amplivasatus) and the oligochaete Tubificoides spp.. These taxa are generally accompanied by Nephtys hombergii whilst the polychaetes Capitella capitata and Melinna palmata may also occur in high numbers in some areas. Other members of the cirratulid polychaete group e.g. Caulleriella zetlandica and Tharyx spp. may also occur in high numbers, sometimes replacing Aphelochaeta marioni as the dominant polychaete. However, there is still inconsistency in the identification of the cirratulid group which is further 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 (Information taken from Connor et al., 2004).

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

Search on:

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 marioni varies 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 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).
  • 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?

    Sensitivity characteristics of the habitat and relevant characteristic species

    SS.SMu.SMuVS.AphTubi is an infralittoral biotope occurring in sheltered, very sheltered and extremely sheltered areas with moderately strong and weak tidal streams (Connor et al., 2004). The biotope occurs in cohesive mud and sandy mud, possibly with shell debris and stones, in full and variable salinities. The biotope is dominated by the polychaete Aphelochaeta marioni (or other Aphelochaeta species e.g. Aphelochaeta amplivasatus) and oligochaete Tubificoides spp.. These taxa are generally accompanied by high numbers of other polychaetes.

    SS.SMu.SMuVS.NhomTubi is a variable salinity biotope, occurring in soft infralittoral mud and sandy mud characterized by the polychaete Nephtys hombergii and oligochaetes of the genus Tubificoides (Connor et al., 2004). Other characterizing species that may be present are the polychaetes Streblospio shrubsolii and Aphelochaeta marioni, and the cumacean Diastylis rathkei typica.

    SS.SMu.SMuVS.NhomTubi is found in areas of silt deposition in soft and sandy muds but may not form a stable habitat. It may be found adjacent to SS.SMu.SMuVS.AphTubi, separated by the abundance of Aphelochaeta marioni and its more cohesive sediments. SS.SMu.SMuVS.AphTubi and SS.SMu.SMuVS.NhomTubi and therefore assessed here as a group with the relevant physical and biological differences highlighted where necessary.

    SS.SMu.SMuVS.AphTubi and SS.SMu.SMuVS.NhomTubi are dominated by the polychaete Aphelochaeta marioni and oligochaete Tubificoides spp., and Nephtys hombergii and oligochaete Tubificoides spp., respectively. These species are, therefore, the focus of this assessment. Other taxa present contribute to species richness and diversity but are not considered important characterizing, defining or structuring species and are not considered within the assessment. More information on these species can be found in other biotope assessments available on this website.

    Resilience and recovery rates of habitat

    Aphelochaeta marioni is a thin, thread like, segmented worm, typically between 2 and 3.5 cm in length, although individuals can reach 10 cm in length (Rayment, 2007a). It lives buried in the upper 4 cm of soft sediments, with the smaller animals nearer the surface. Aphelochaeta marioni can live up to 2-3 years and its lifecycle varies according to environmental conditions (Rayment, 2007a). In Stonehouse Pool, Plymouth Sound, 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). Laboratory observation reported spawning occurring at night, with females that rose up into the water column with their tail end in the burrow, and shed eggs within a few seconds that sank to form puddles on the sediment (Farke, 1979). Fertilization was not observed, probably because the male does not leave the burrow. The embryos developed lecithotrophically and hatched in about 10 days (Farke, 1979). The newly hatched juveniles were ca 0.25 mm in length and immediately dug into the sediment. Where the sediment depth was not sufficient for digging, the juveniles swam or crawled in search of a suitable substratum (Farke, 1979). In the laboratory, juvenile mortality was high (ca 10% per month) and most animals survived for less than a year (Farke, 1979). In the Wadden Sea, the majority of the cohort reached maturity and spawned at the end of their first year, although some slower developers did not spawn until the end of their second year (Farke, 1979). However, the population of Aphelochaeta marioni in Stonehouse Pool spawned for the first time at the end of their second year (Gibbs, 1971). There was no evidence of a major post-spawning mortality and it was suggested that individuals may survive to spawn over several years. Gibbs (1971) found that the number of eggs laid varied from 24-539 (mean=197) and was correlated with the female's number of genital segments and, hence, female size and age.

    Usually for oligochaetes, fertilization is internal and relatively few large eggs are shed directly into a cocoon that is secreted by the worm (Giere & Pfannkuche, 1982). Asexual reproduction is possible in some species by spontaneous fission (Giere & Pfannkuche, 1982). The naid oligochaete Panais litoralis can produce asexually producing clones, the rapid rate of increase (18 times population abundance in 3 months, Gillett et al., 2007) allows this species (which is sensitive to high temperatures, hypoxia and is exposed to predation due to shallow burial) to repopulate rapidly when conditions are favourable.  However, few Tubificidae and Enchytraeidae produce asexually (Giere & Pfannkuche, 1982).

    Tubificid populations tend to be large and to be constant throughout the year, although some studies have noticed seasonal variations (Giere & Pfannkuche, 1982). Many species, including Tubificoides benedii have a two-year reproductive cycle and only part of the population reproduces each season (Giere & Pfannkuche, 1982). Populations of Tubificoides benedii in the Fourth Estuary have not demonstrated clear seasonality in recruitment (Bagheri & McLusky, 1982), although mature Tubificoides benedii (as Peloscolex benedeni) in the Thames Estuary were reported to occur in December with a maximum in late February (Hunter & Arthur, 1978), breeding worms increased from April and maximum cocoon deposition was observed in July (Hunter & Arthur, 1978). Tubificids exhibit many of the traits of opportunistic species. It is dominant, often reaching huge population densities in coastal areas that are enriched in organic matter and is often described as an ‘opportunist’ species adapted to rapid environmental fluctuations and stress (Giere, 2006; Bagheri & McLusky, 1982). However, unlike other opportunist species it has a long-life span (a few years, Giere, 2006), a prolonged reproductive period from reaching maturity to maximum cocoon deposition and exhibits internal fertilization with brooding rather than pelagic dispersal. These factors mean that recolonization is slower than for some opportunistic species such as Capitella capitata and nematodes which may be present in similar habitats.

    Bolam & Whomersley (2003) observed faunal recolonization of fine sediments placed on saltmarsh as a beneficial use and disposal of fine grained dredged sediments. They found that tubificid oligochaetes began colonizing sediments from the first week following a beneficial use scheme involving the placement of fine-grained dredged material on a saltmarsh in south east England. The abundance of Tubificoides benedii recovered slowly in the recharge stations and required 18 months to match reference sites and those in the recharge stations prior to placement of sediments. The results indicate that some post-juvenile immigration is possible and that an in situ recovery of abundance is likely to require more than 1 year.  Rapid recolonization has also been observed in the tubificid oligochaete  Baltidrilus costata (Tubifex costatus) appeared  in upper sediment layers in experimentally defaunated patches (4 m2)  after 3 weeks (Gamenick et al., 1996).

    Dittman et al. (1999) observed that Nephtys hombergii was amongst the macrofauna that colonized experimentally disturbed tidal flats within two weeks of the disturbance that caused defaunation of the sediment. However, if sediment is damaged, recovery is likely to be slower, for instance, Nephtys hombergii abundance was reduced by 50% in areas where tractor towed cockle harvesting was undertaken on experimental plots in Burry Inlet, South Wales, and had not recovered after 86 days (Ferns et al., 2000). Nephtys hombergii has a 3-10 year lifespan, and reaches maturity at 1-2 years. In the Tyne Estuary, spawning of Nephtys hombergii occurred in May and September, whilst in Southampton Water the species spawned throughout the year with peaks in July and November (Wilson, 1936; Oyenekan, 1986). In Århus Bay, Denmark, Nephtys hombergii spawned in August and September, but a decrease in the number of individuals bearing gametes in May and June suggested that at least part of the population spawned in early summer (Fallesen & Jørgensen, 1991). Two larval cohorts were observed from Nephtys hombergii in Arachon Bay, France within one year (Mathivat-Lallier & Cazaux, 1991). Post-larvae settled directly in the intertidal area and again two distinct waves of recruitment were observed. These traits suggest the species is likely to require longer to recover than more opportunistic species.

    Resilience assessment. The biotopes are dominated by fast growing opportunistic polychaetes and are likely to reach maturity within one year of space becoming available. Seasonal changes may, however, occur in the abundance of the fauna due to seasonal recruitment processes. For example, variation in abundance is very pronounced in the polychaete Aphelochaeta marioni. In the Wadden Sea, peak abundance occurred in January (71,200 individuals per m²) and minimum abundance occurred in July (22,500 individuals per m²), following maximum spawning activity between May and July (Farke, 1979). In general, there was little information found for Tubificoides benedii, but, taking into consideration the information above (particularly Bolam & Whomersley, 2003), this review considers that the recoverability of this species is generally high. Recovery of Nephtys hombergii is likely to occur over 1-2 years. Removal of the characterizing species Aphelochaeta, Tubificoides spp. and Nephtys hombergii would likely result in the biotopes being lost and reclassified. Where a disturbance event removes part of the community (resistance High, Medium or Low) and recruitment of the characterizing species is possible by adult migration or recruitment from the remaining members of the community, resilience is likely to be High. However, Aphelochaeta marioni has no pelagic phase in its lifecycle, and dispersal is limited to the slow burrowing of the adults and juveniles (Farke, 1979). So where the community is severely reduced (resistance None), recruitment is likely to depend on dispersal by hydrodynamic conditions. The low energy conditions experienced by SS.SMu.SMuVS.AphTubi may limit immediate recolonization. However, as long as the substratum nature of the biotope remains suitable for the settlement of recruits, the community is likely to reach maturity within 2-10 years, so resilience of SS.SMu.SMuVS.AphTubi is likely to be Medium. On the other hand, Aphelochaeta is less abundant in SS.SMu.SMuVS.NhomTubi and recovery of the biotope from severe defaunation (resistance None) will depend on successful recruitment of Nephtys hombergii and Tubificoides spp., which will be aided by more energetic conditions. Thus, recovery of SS.SMu.SMuVS.NhomTubi is still likely to be High.

    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 prior to impact.  This does not necessarily mean that every component species has returned to its prior condition, abundance or extent but that the relevant functional components are present and the habitat is structurally and functionally recognizable as the initial habitat of interest. It should be noted that the recovery rates are only indicative of the recovery potential.

    Hydrological Pressures

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

    Aphelochaeta marioni is distributed over a wide temperature range. It has been recorded from the Mediterranean Sea and Indian Ocean (Farke, 1979). Therefore, the species must be capable of tolerating higher temperatures than it experiences in Northern Europe. For example, Covazzi Harriague et al. (2007) reported Aphelochate marioni occurring in the Rapallo Harbour (Ligurian Sea, NW Mediterranean) at 24°C. Furthermore, Aphelochaeta marioni lives infaunally and so is likely to be insulated from rapid temperature change. An increase in temperature would be expected to cause some physiological stress but no mortality.

    Deeper burrowing oligochaetes are protected from fluctuations in temperature by the overlaying sediments which dampen changes if poorly drained (Giere & Pfannkuche, 1982).  Bamber & Spencer (1984) observed that Tubificoides were dominant species in an area affected by thermal discharge in the River Medway estuary. Sediments were exposed to the passage of a temperature front of approximately 10oC between heated effluent and estuarine waters during the tidal cycles. Tubificoides benedii has increased in abundance in mudflat habitats in Jade Bay, North Sea between 1930 and 2009 (Schueckel & Kroencke, 2013). Climate warming as well as decreasing nutrient loads and species introductions have occurred in the region since the 1970s, suggesting the species may adapt to temperature increases at the benchmark level. Tubificoides benedii is considered an opportunist that is adapted to rapid environmental fluctuations and harsh conditions in estuaries (Gogina et al., 2010), suggesting resistance would be high to this pressure at the benchmark level.

    Found from the northern Atlantic, from such areas as the Barents Sea, the Baltic and the North Sea, to the Mediterranean, Nephtys hombergii has been reported from as far south as South Africa, suggesting the species can tolerate a 5°C increase in temperature in the UK and Irish coasts. Records are limited but Emery & Stevensen (1957) found that Nephtys hombergii could withstand summer temperatures of 30-35°C.

    Environmental factors, such as temperature, day length, and tidal or lunar cycles, have been implicated in the timing of spawning of Nephtyidae, in particular the spring tide phase of the lunar cycle (Bentley et al., 1984). In the Tyne Estuary, spawning of Nephtys hombergii occurred in May and September, whilst in Southampton Water the species spawned throughout the year with peaks in July and November (Wilson, 1936; Oyenekan, 1986). In Århus Bay, Denmark, Nephtys hombergii spawned in August and September, but a decrease in the number of individuals bearing gametes in May and June suggested that at least part of the population spawned in early summer (Fallesen & Jørgensen, 1991). A 5°C increase in temperature for one month period, or 2°C for one year is likely to impact the timing of reproduction in these areas. A combination of environmental factors appears to influence timing (in particular spring tides), therefore, temperature may not be the most significant cue but evidence is limited to identify the significance of temperature in relation to other factors (or cues).

    Sensitivity assessment. Typical surface water temperatures around the UK coast vary seasonally from 4-19°C (Huthnance, 2010). Most organisms in the biotopes are distributed to the north and south of Britain and Ireland and unlikely to be affected adversely by long-term temperature change. In addition, subtidal and especially infaunal species are likely to be protected from acute temperature change. Furthermore, the dominance of Tubificoides spp. when exposed to a heated effluent suggests that this genus would be highly resistant to an increase in temperature at the pressure benchmark. Resistance and resilience of the biotopes are therefore assessed as High and the biotopes are judged as Not Sensitive.

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

    Aphelochaeta marioni is distributed over a wide temperature range. It has been recorded from the western Baltic Sea, south Atlantic Ocean and North Sea (Farke, 1979). Therefore the species must be capable of tolerating low temperatures. Aphelochaeta marioni lives buried in sediment and is therefore well insulated from decreases in temperature. In the Wadden Sea, the population was apparently unaffected by a short period of severe frost in I973 (Farke, 1979). Kędra et al. (2010) reported Aphelochaete marioni occurring in the Svalbard Archipelago where temperatures below zero may be experienced in the winter. A decrease in temperature would be likely to cause some physiological stress but no mortality.

    Most littoral oligochaetes, including tubificids, can survive freezing temperatures and can survive in frozen sediments (Giere & Pfannkuche, 1982). Tubificoides benedii (studied as Peloscolex benedeni) recovered after being frozen for several tides in a mudflat (Linke, 1939).

    Nephtys hombergii are found as far north as the Barents Sea, and would be expected to be resistant to a 5°C decrease in temperature for one month period, or 2°C for one year. Environmental factors, such as temperature, day-length, and tidal or lunar cycles, have been implicated in the timing of spawning of the Nephtyidae, in particular the spring tide phase of the lunar cycle (Bentley et al., 1984). Olive et al. (1997) found that relative spawning success in a North Sea (Newcastle on Tyne) population of Nephtys hombergii was positively correlated with winter sea and air temperatures. This suggests a 5°C decrease in temperature for a one month period, occurring in winter, or 2°C for one year are likely to impact spawning success.

    Sensitivity assessment. Typical surface water temperatures around the UK coast vary seasonally from 4-19°C (Huthnance, 2010). Based on the evidence presented, Aphelochaeta marioni, Tubificoides benedii and Nephtys hombergii 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. Temperature may act as a spawning cue and an acute or chronic decrease may result in some delay in spawning, however, this is not considered to impact the adult populations and may be compensated by later spawning events. Resistance and resilience are therefore assessed as High and the biotopes judged as Not Sensitive.

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

    Environmental fluctuations in salinity are only likely to affect the surface of the sediment, and not deeper buried organisms, since the interstitial or burrow water is less affected. However, under longer term or permanent increase in salinity, sediment waters would also adjust.

    Populations of Aphelochaeta marioni inhabit the open coast where seawater is at full salinity. Covazzi Harriague et al. (2007) reported Aphelochaeta marioni in the Rapallo Harbour (Ligurian Sea, NW Mediterranean) at salinities above 38 psu. Farke (1979) studied the effects of changing salinity on Aphelochaeta marioni (studied as Tharyx marioni) in a microsystem in the laboratory. Over several weeks, the salinity in the microsystem was increased from 25-40 psu and no adverse reaction was noted. However, when individuals were removed from the sediment and displaced to a new habitat, they only dug into their new substratum if the salinities in the two habitats were similar. If the salinities differed by 3-5 psu, the worms carried out random digging movements, failed to penetrate the sediment and died at the substratum surface after a few hours. This would suggest that Aphelochaeta marioni can tolerate salinity changes when living infaunally but may not resist an increase in salinity when removed from its habitat.

    Oligochaete dominated biotopes are recorded from a range of salinity regimes from full (LS.LSa.MoSa.Ol; LS.LSa.MoSa.Ol.FS), variable (SS.SMu.SMuVS.CapTubi) to low (SS.SMu.SMuVS.LhofTtub) habitats (Connor et al., 2004). The species characterizing these biotopes are likely to vary. Giere & Pfannkuche (1982) identified how species change over a hypothetical salinity gradient with marine stenohaline species present at full salinities replaced by more euryhaline oligochaete species including Tubificoides benedii and Tubificoides pseudogaster. Studies in the Rhine delta have found that Tubificoides benedii is tolerant of a range of salinities (Verdonschot et al., 1982).

    Nephtys hombergii is considered to be a brackish water species (Barnes, 1994), but where the species occurs in open coastal locations the species would have to tolerate salinities of 25 psu and above. Within a few months of the closure of a dam across the Krammer-Volkerak estuary in the Netherlands, Wolff (1971b) observed that species with pelagic larvae or a free-swimming phase, including Nephtys hombergii expanded rapidly with a concomitant increase of salinity to 9-15 psu everywhere. Prior to the closure of the dam, the estuary demonstrated characteristics of a typical 'salt-wedge' estuary with a salinity gradient from 0.3 to 15 psu. Hence, Nephtys hombergii is likely to survive increases in salinity within estuarine environments. Nephtys hombergii may still be found in fully marine locations but, may be competitively inferior to other species of Nephtyidae (e.g. Nephtys ciliata and Nephtys hystricis) and occur in lower densities. An increase to fully marine (30-40 ‰) would therefore be likely to lead to a reduction in density of Nephtys hombergii.

    Sensitivity assessment. Based on that all species occur in greater salinities than those that occur in these biotopes, the biological assemblages associated with the biotopes are considered to have High resistance and High resilience and is therefore considered to be Not Sensitive to an increase in salinity at the pressure benchmark level.

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

    Environmental fluctuations in salinity are only likely to affect the surface of the sediment, and not deeper buried organisms, since the interstitial or burrow water is less affected. However, under longer term or permanent increase in salinity, sediment waters would be expected to also adjust.

    Aphelochaeta marioni thrives in estuaries and is, therefore, likely to be tolerant of decreases in salinity. It 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 freshwater discharge from rivers is high (Farke, 1979). In the Severn Estuary, Aphelochaeta marioni (studied as Tharyx marioni) characterized the faunal assemblage of very poorly oxygenated, poorly sorted mud with relatively high interstitial salinity (Broom et al., 1991).

    Nephtys hombergii is considered to be a brackish water species, and has been reported to extend in to estuarine locations where salinity is less than 18 psu (Barnes, 1994). Clark & Haderlie (1960) found Nephtys hombergii in the Bristol Channel at salinities between 15.9 psu and 25.1 psu. If the salinity were to become intolerable to the polychaete, it is likely that as a mobile species, able to both swim and burrow, Nephtys hombergii would avoid the change in salinity by moving away and localized densities would decline.

    In Ria de Averio, western Portugal, Tubificoides benedii characterized communities in estuarine sample sites further upstream with lower salinity, suggesting a high resistance to a decrease in salinity (Rodrigues et al., 2011).

    Overall, the important characterizing species are likely to tolerate a short-term change in salinity from e.g. variable to low salinity and a long-term change from variable to reduced salinity. The species richness of the biotopes may decline but the biotopes will probably not be adversely affected.

    Sensitivity assessment. Records indicate SS.SMu.SMuVS.AphTubi occurs in areas of full (30-35 ppt) and variable (18-35 ppt) salinities, whereas SS.SMu.SMuVS.NhomTubi occurs in areas of variable (18-35 ppt) salinity (Connor et al., 2004). Nephtys hombergii is possibly the most sensitive to the lowest salinity levels within the ‘low’ salinity category, although as a mobile species, it will be resistant through being able to move lower down the shore or away from freshwater run off. The remaining characterizing species Aphelochaeta marioni and Tubificoides spp. occur in estuarine areas experiencing low salinities, so are therefore likely to resist a decrease in salinity at the pressure benchmark level. Resistance of the biotopes is therefore assessed as High and resilience as High (by default), and the biotopes are considered Not Sensitive to a decrease in salinity at the pressure benchmark level.

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

    The hydrographic regime is an important structuring factor in sedimentary habitats. An increase in water flow rate is not likely to affect Nephtys hombergii, Aphelochaeta marioni and Tubificoides spp. and other characterizing species as they live infaunally. The most damaging effect of increased flow rate could be the erosion of the substratum as this could eventually lead to loss of the habitat. Orvain et al. (2007) investigated the spatio-temporal variations in intertidal mudflat erodibility in western France and suggested a potential link between Polychaeta and bed erodibility given the high polychaete abundances observed in the study.

    Increased water flow rates is likely to change the sediment characteristics in which the species live, primarily by resuspending and preventing deposition of finer particles (Hiscock, 1983). The characterizing species prefer habitats with silty/muddy substrata which would not occur in very strong tidal streams. Coarser sediments are likely to remain in areas of strongest flow velocity (where finer particles have been resuspended) (Coates et al., 2014). Species such as Tubificoides benedii and other opportunist polychaetes that tolerate coarser particle size will possibly increase in abundance. Aphelochaeta marioni has been recorded in the Wadden Sea with flow rates up to 0.45 m/s (Compton et al., 2013), and in the Westerscheld estuary (SW Netherlands) where spring current velocities vary between 0.1-0.58 m/s (Van Colen et al., 2010a).

    Additionally, the consequent lack of deposition of particulate matter at the sediment surface would reduce food availability. Decreased water movement would result in increased deposition of suspended sediment (Hiscock, 1983). An increased rate of siltation resulting from a decrease in water flow may result in an increase in food availability for the characterizing species and therefore growth and reproduction may be enhanced, but only if food was previously limiting.

    Sensitivity assessment. Sand particles are most easily eroded and likely to be eroded at about 0.20 m/s (based on Hjulström-Sundborg diagram, Sundborg, 1956). Although having a smaller grain size than sand, clays and silts require greater critical erosion velocities because of their cohesiveness. The biotopes occur most often in weak, moderately strong and strong tidal streams (Connor et al., 2004), suggesting the species characterizing the biotopes can adapt to a range of tidal currents, aided by each species burrowing traits. A change in water flow rate at the pressure benchmark level of 0.1-0.2 m/s is considered to fall within the range of flow speeds experienced by populations in the middle of their range. Resistance and resilience are, therefore, assessed as High and the biotopes considered Not Sensitive to a change in water flow at the pressure benchmark level.

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

    The biotopes occur in the infralittoral so that only the upper extent of shallow examples are likely to be emersed on extreme low tides. All characterizing species would probably survive an increase in emergence. However, the species can only feed when immersed and therefore likely to experience reduced feeding opportunities. Over the course of a year, the resultant energetic cost is likely to cause some mortality. In addition, increased emergence is likely to increase the vulnerability to predation from shore birds. A decrease in emergence is likely to allow the biotopes to extend their upper limits, where suitable substrata exist.

    Opportunistic, deposit feeding polychaetes and oligochaetes, such as Aphelochaeta marioni and Tubificoides benedii are likely to tolerate stressful conditions, and often outcompete more sensitive species in intertidal environments due to greater tolerances (Gogina et al., 2010). Nephtys hombergii is sufficiently mobile to rapidly burrow and seek damper substrates during periods when emergence increases. For instance, Vader (1964) observed that the worm relocates throughout the tidal cycle.

    For instance, Tubificoides benedii is capable of penetrating the substrate to depths of 10 cm, shows resistance to hypoxia and is often typified as an ‘opportunist’ that is adapted to the rapid environmental fluctuations and harsh conditions in estuaries (Gogina et al., 2010). Highest abundances were predicted by Gogina et al. (2010) to be related to depth with an optimum of 10 m to 20 m. Further case studies were not returned by literature searches on Tubificoides benedii. The studies returned by searches suggest abundance may be limited by a decrease in high water level or a change in time (increase) where substrate is not covered by the sea. An increase in the time the biotopes are covered by the sea is likely to result in increased abundance of Tubificoides benedii.

    Sensitivity assessment. Some mortality of the characterizing species is likely to occur because of emergence regime changes. Resistance is therefore assessed as Medium and resilience is likely to be High, so the biotopes are considered to have Low sensitivity to emergence regime changes at the pressure benchmark level.

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

    Strong wave action is likely to cause damage or withdrawal of delicate feeding and respiration structures resulting in loss of feeding opportunities and compromised growth for the characterizing species. Decreases in wave exposure may influence the supply of particulate matter because wave action may have an important role in resuspending the sediment. Furthermore, Aphelochaeta marioni characteristically inhabits soft sediments in sheltered areas (Broom et al., 1991), so individuals may be damaged or dislodged by scouring from sand and gravel mobilized by increased wave action.

    Disturbance of sediment by waves may reduce oligochaete abundance (Giere, 1977) and oligochaetes may be absent from very wave exposed shores (Giere & Pfannkuche, 1982). 

    Nephtys hombergii lives infaunally but may sometimes partially emerge to seek and capture food but does not present a significant surface area to wave action to sustain physical damage. Clark & Haderlie (1960) and Clark et al. (1962) suggested that strong wave action limited the distribution of Nephtys hombergii. Increased wave action for long durations (e.g. 1 year) may ultimately change the nature of the substratum that the polychaete inhabits and its distribution may consequently alter.

    Sensitivity assessment. Hydrographic regimes are an important structuring factor in sedimentary habitats, and an increase in wave exposure could result in fine sediments being eroded (Hiscock, 1983), resulting in the likely reduction of the habitat and a decrease in food availability. Some erosion will occur naturally and storm events may be more significant in loss and damage of the substratum than changes in wave height at the pressure benchmark. Limited zoobenthic biomass has been recorded in areas exposed to strong currents and wave action (Beukema, 2002), limiting food availability to species such as Nephtys hombergii, however, impacts from this pressure at the benchmark level may be low for the biotopes, as they are limited to sheltered locations. Increases in wave action may therefore remain within the limits of the species tolerances but factors such as sediment redistribution may alter the physical biotopes. The biotopes are found in moderately exposed, sheltered, very sheltered and extremely sheltered sites (Connor et al., 2004) and a change at the benchmark level is likely to fall within the range experienced by the mid-range examples. The biotopes are therefore considered to have High resistance to changes at the pressure benchmark, where these do not lead to increased erosion of the substratum. Resilience is therefore assessed as High and the biotopes are considered to be Not Sensitive at the pressure benchmark.

    Chemical Pressures

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

    Evidence suggests that polychaetes are fairly resistant to the effects of heavy metals (Bryan, 1984). Aphelochaeta marioni is tolerant of heavy metal contamination occurring in the heavily polluted Restronguet Creek (Bryan & Gibbs, 1983) and it is also an accumulator of arsenic (Gibbs et al., 1983). Nephtys hombergii is also recorded in Restronguet Creek. Rygg (1985) classified Tubificoides spp. as highly tolerant species, common at the most copper polluted stations (>200 mg/kg) in Norwegian fjords.

    Sensitivity assessment. Taking account of the variable salinity conditions that affect these biotopes (in general, for estuarine animals, heavy metal toxicity increases as salinity decreases and temperature increases: McLusky et al., 1986), it seems possible that some polychaete species, at least in the biotopes, might be adversely affected by high contamination by heavy metals. However, the biotopes are considered Not Sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

    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

    Cirratulids seem to be mostly immune to oil spills, probably because their feeding tentacles are protected by a heavy secretion of mucus (Suchanek, 1993). This is supported by observations of Aphelochaeta marioni following the Amoco Cadiz oil spill in March, 1978 (Dauvin, 1982, 2000). Prior to the spill, Aphelochaeta marioni (studied as Tharyx marioni) was present in very low numbers in the Bay of Morlaix, western English Channel. Following the spill, the level of hydrocarbons in the sediment increased from 10 mg/kg dry sediment to 1443 mg/kg dry sediment 6 months afterwards. In the same period, Aphelochaeta marioni increased in abundance to a mean of 76 individuals/m2, which placed it among the top five dominant species in the faunal assemblage. It was suggested that the population explosion occurred due to the increased food availability because of accumulation of organic matter resulting from high mortality of browsers. Six years later, abundance of Aphelochaeta marioni began to fall away again, accompanied by gradual decontamination of the sediments.

    Tubificoides benedii was found in UK waters near oil refineries as the sole surviving member of the macrofauna. Populations were, however, apparently reduced and the worms were absent from areas of oil discharge and other studies indicate sensitivity to oiling (Giere & Pfannkuche, 1982).

    McLusky (1982) found that petrochemical effluents, including organic solvents and ammonium salts, released from a point source to an estuarine intertidal mudflat of the Forth Estuary, Scotland, caused severe pollution in the immediate vicinity. Beyond 500 m distance, the effluent contributed to an enrichment of the fauna in terms of abundance and biomass similar to that reported by Pearson & Rosenberg (1978) for organic pollution; Nephtys hombergii was found in the area with maximum abundance of species and highest total biomass at 500 m from the discharge. It seems likely that significant hydrocarbon contamination would kill affected populations of the species. On return to prior conditions, recolonization is likely via adult migration and larval settlement.

    Sensitivity assessment. Overall, hydrocarbon contamination is likely to adversely affect some members of the community, and low more tolerant or opportunistic species to increase in abundance, resulting in a reduction in species richness. However, the biotopes are considered Not Sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

    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

    There is little evidence directly relating to the effects of synthetic chemicals on Aphelochaeta marioni. Waldock et al. (1999) reported that the species diversity of polychaete infauna, including Aphelochaeta marioni, in the Crouch Estuary increased in the three years after the use of TBT was banned within the estuary, suggesting that TBT had suppressed their abundance previously. Furthermore, Beaumont et al. (1989) concluded that TBT had a detrimental effect on the larval and/or juvenile stages of infaunal polychaetes.

    No evidence concerning the specific effects of chemical contaminants on Nephtys hombergii was found. Boon et al. (1985) reported that Nephtys species in the North Sea accumulated organochlorines but, based on total sediment analyses, organochlorine concentrations in Nephtys species were not correlated with the concentrations in the (type of) sediment which they inhabited.

    Sensitivity assessment. The biotopes are considered Not Sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

    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 is available on which to assess this pressure.

    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

    The biotopes are considered to be Not Sensitive at the pressure benchmark that assumes compliance with all relevant environmental protection standards.

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

    Connor et al. (1997a) described sediments in which Aphelochaeta marioni is commonly found as usually with a "black anoxic layer close to the sediment surface." Broom et al. (1991) recorded that Aphelochaeta marioni (studied as Tharyx marioni) characterized the faunal assemblage of very poorly oxygenated mud in the Severn Estuary. They found Aphelochaeta marioni to be dominant where the redox potential at 4 cm sediment depth was 56 mV and, therefore, concluded that the species was tolerant of very low oxygen tensions. It is likely that feeding, growth and reproduction would be impaired under sustained low oxygen conditions.

    Oligochaetes, such as Tubificoides spp. vary in their tolerance of hypoxia and associated high sulphide levels. Most enchytraaids and naidids are sensitive to hydrogen sulphide and hypoxia while tubificids are often more resistant (Giere, 2006). Tubificoides benedii has a high capacity to tolerate anoxic conditions, its extreme oxygen tolerance is based on an unusually low respiration rate (Giere et al., 1999). Respiration rates of Tubificoides benedii measured at various oxygen concentrations showed that aerobic respiration is maintained even at very low oxygen concentrations (Giere et al., 1999). Birtwell & Arthur (1980) showed that Tubificoides benedii could tolerate anoxia in the Thames Estuary (LT50 = 58.8 hours at 20oC, 26.6 hours at 25oC  and 17.8 hours at 30oC in experiments with worms acclimated to 20oC.)

    Nephtys hombergii inhabits intertidal areas where resistance to low oxygen is needed and sulphide levels can reach up to 1 mM (Giere, 1992; Thierman et al., 1996). As with other characterizing polychaete species, Nephtys hombergii exhibits the ability to switch from aerobic to anerobic respiration, which provides some protection from the toxic effects of sulphide. Nephtys hombergii has adapted to such conditions by utilizing several strategies. Arndt & Schiedek (1997) found Nephtys hombergii to have a remarkably high content of phosphagen (phosphoglycocyamine), which is the primary energy source during periods of environmental stress. With increasing hypoxia, energy is also provided via anaerobic glycolysis, with strombine as the main end-product. Energy production via the succinate pathway becomes important only under severe hypoxia, suggesting a biphasic response to low oxygen conditions which probably is related to the polychaete's mode of life. The presence of sulphide resulted in a higher anaerobic energy flux and a more pronounced energy production via glycolysis than in anoxia alone. Nevertheless, after sulphide exposure under anaerobic conditions of <24 h, Arndt & Schiedek (1997) observed Nephtys hombergii to recover completely. Although Nephtys hombergii appears to be well adapted to a habitat with short-term fluctuations in oxygen and appearance of hydrogen sulphide, its high energy demand as a predator renders it likely to limit its survival in an environment with longer lasting anoxia and concomitant sulphide exposure. For instance, Fallesen & Jørgensen (1991) recorded Nephtys hombergii in localities in Århus Bay, Denmark, where oxygen concentrations were permanently or regularly low, but in the late summer of 1982 a severe oxygen deficiency killed populations of Nephtys species (Nephtys hombergii and Nephtys ciliata) in the lower part of the bay. However, Nephtys hombergii recolonized the affected area by the end of autumn the same year. Alheit (1978) reported a LC50 at 8°C of 23 days for Nephtys hombergii maintained under anaerobic conditions.

    Nephtys hombergii have tolerated extreme hypoxia, leaving the sediment only after 11 days (Nilsson & Rosenberg, 1994). Nephtys hombergii in artificially created anoxic conditions were shown to survive for at least 5 days (Schöttler, 1982) and do not switch from aerobic to anerobic metabolic pathways until oxygen saturation decreases <12% (Schöttler, 1982).

    Sensitivity assessment. The characterizing species are likely to only be affected by severe deoxygenation episodes. Resistance to deoxygenation at the pressure benchmark level is likely to be High. Resilience of the biotopes is likely to also be High and the biotopes are therefore considered Not Sensitive to exposure to 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

    This pressure relates to increased levels of nitrogen, phosphorus and silicon in the marine environment compared to background concentrations.  The benchmark is set at compliance with WFD criteria for good status, based on nitrogen concentration (UKTAG, 2014). The biotopes are, therefore, considered Not Sensitive at the pressure benchmark that assumes compliance with good status as defined by the WFD.

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

    Kędra et al. (2010) reported Aphelochaeta mariori that occurred in the Hornsundfjord, Svalbard, where primary production has been recorded as 120 gC/m2/yr. Covazzi Harriague et al. (2007) reported Aphelochaeta mariori at sites with organic matter sedimentation as high as 359 mg/m2/hr in the Ligurian Sea, Italy. Furthermore, Markert et al. (2010) compared macrofaunal communities in the Wadden Sea in reefs dominated by Mytilus edulis and Cassostrea gigas and found Aphelochaeta mariori as a dominate species throughout the study site, which suggested the species was unlikely to be affected by organic enrichment.

    Tubificoides benedii is very tolerant of high levels of organic enrichment and often dominate sediments where sewage has been discharged or other forms of organic enrichment have occurred (Pearson & Rosenberg, 1978; Gray, 1971; McLusky et al., 1980). Their tolerance for organic enrichment is attributed to their adaptation to live in and feed on enriched organic deposits (Pearson & Rosenberg, 1978), and their high population densities in such areas is enhanced by the lack of predation and competition. Tubificoides benedii are abundant in mussel beds (mussel relaying may be the source of smothering) which has been attributed to their tolerance of organically rich deoxygenated sediment (Commito & Boncavage, 1989). Tubificoides benedii has also been found in elevated abundances in areas of organic enrichment around fish farms (Haskoning, 2006).

    The infaunal habit of Nephtys hombergii and its ability to burrow relatively rapidly through, and into the substratum are likely to aid the species in its avoidance of unsuitable conditions. Nephtys spp. were present in organically enriched sediments  along the Catalan Spanish coast (Cardell et al., 1998), suggesting the species is likely to be resistant to some organic enrichment.

    Sensitivity assessment. The evidence presented suggest the characterizing species may not be affected by organic enrichment at the benchmark level. Resistance and resilience are therefore assessed as High and the biotopes are considered Not Sensitive to organic enrichment (deposit of 100 gC/m2/yr).

    Physical Pressures

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

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

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

    If the cohesive mud and sandy mud that characterize these biotopes were replaced with rock substrata, this would represent a fundamental change to the physical character of the biotopes. The characterizing species would no longer be supported and the biotopes would be lost and/or reclassified.

    Sensitivity assessment. Resistance to the pressure is considered None and resilience Very Low. 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: Medium
    A: Medium
    C: High
    Q: High
    A: High
    C: High
    Q: Medium
    A: Medium
    C: High

    Aphelochaeta marioni has been recorded from a variety of different sediment types. In the intertidal area of the Wadden Sea, it achieved highest abundance where the sediment fraction smaller than 0.04 mm diameter was greater than 10% of the total sediment (Farke, 1979). Tubificoides benedii (studied as Peloscolex benedeni) is found in a range of substratum types from sandy mixed habitats, fine sands and coarse sands (Giere & Pfannkuche, 1982). High densities of Nephtys hombergii were found in substrata of 0.3% particles >0.25 mm and 5.8% <0.125 mm in diameter, but the worm tolerated up to 3.8% 0.25 mm and 2.2-15.9% <0.125 mm (Clark et al., 1962). Nephtys hombergii may be found in higher densities in muddy environments and this tends to isolate it from Nephtys cirrosa, which is characteristic of cleaner, fairly coarse sand. Degraer et al. (2006) summarized that the higher the medium grain size of the sediment, the lower the relative occurrence of Nephtys hombergii and in grain sizes over 0.5 mm the species was absent in the Belgium part of the North Sea.

    Sensitivity assessment. The characterizing species of these biotopes occur in a range of substrata and are likely to resist a change in sediment type by 1 Folk class from, for example, sandy mud to muddy sand (based on the Long, 2006 simplification). However, this would probably represent a fundamental change in the character of the biotopes, and a change in the abundance of the characteristic species, resulting in the loss and/or reclassification of the biotopes. Resistance is therefore assessed as None and resilience as Very Low given the permanent nature of the pressure and the biotopes are considered to have High sensitivity to a change in seabed type by one Folk class.

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

    Sedimentary communities are likely to be highly intolerant of substratum removal, which will lead to partial or complete defaunation, and will expose underlying sediment which may be anoxic and/or of a different character or bedrock and lead to changes in the topography of the area (Dernie et al., 2003). Any remaining species, given their new position at the sediment/water interface, may be exposed to conditions to which they are not suited. Removal of 30 cm of surface sediment will remove the infaunal community and other species present in the biotopes. For instance, Aphelochaeta marioni lives buried in soft sediments with the majority of individuals found in the upper 4 cm of the sediment (Rayment, 2007a). Removal of the substratum to 30 cm would result in the loss of the characterizing species. Recovery of the biological assemblage may take place before the original topography is restored, if the exposed, underlying sediments are similar to those that were removed. Hydrodynamics and sedimentology (mobility and supply) influence the recovery of soft sediment habitats (Van Hoey et al., 2008).

    Sensitivity assessment. Extraction of 30 cm of sediment will remove the characterizing biological component of the biotopes. Resistance of the biotopes is assessed as None. Resilience of SS.SMu.SMuVS.AphTubi is likely to be Medium given the low energy environment where the biotope occurs, whereas resilience of SS.SMu.SMuVS.NhomTubi is likely to be High. Both biotopes sensitivity are therefore assessed as Medium.

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

    Aphelochaeta marioni is a soft bodied organism which exposes its palps and cirri at the surface while feeding (Rayment, 2007a). The species lives infaunally in soft sediment, usually within a few centimetres of the sediment surface. Physical disturbance, such as dredging or dragging an anchor, would be likely to penetrate the upper few centimetres of the sediment and cause physical damage to Aphelochaeta marioni. De Biasi & Pacciardi (2008) compared macrobenthic communities in a commercial fishing ground exploited by otter trawling with an area closed to fishing for over 10 years in the Adriatic Sea. The authors found that polychaetes, including Aphelochaeta spp. were among the species dominating the disturbed areas, which is likely to result from the ability of the species to recolonize disturbed areas rapidly, rather than indicate that the polychaetes are resistant to disturbance of the seabed surface.

    Tubificoides benedii can be relatively deeply buried and could avoid direct exposure to abrasion although sediment disturbance and compaction could damage these soft-bodied species, and oligochaetes in general are not found in high abundances in sediments with high levels of disturbance from wave action. Experimental studies on crab-tiling impacts have found that densities of Tubificoides benedii and Tubificoides pseudogaster were higher in non-trampled plots (Sheehan et al., 2010), indicating that these oligochaetes have some sensitivity to trampling. Whomersley et al. (2010) conducted experimental raking on intertidal mudflats at two sites (Creeksea Crouch Estuary England, and Blackness lower Forth Estuary, Scotland), where Tubificoides benedii were dominant species. For each treatment, 1 m2 plots were raked twice to a depth of 4 cm (using a garden rake). Plots were subject to either low intensity treatments (raking every four weeks) or high (raking every two weeks). The experiment was carried out for 10 months at Creeksea and a year at Blackness. The high and low raking treatments appeared to have little effect on Tubificoides benedii (Whomersley et al., 2010).

    Collie et al. (2000) found that Nephtys hombergii displayed a negative effect on abundance as a result of fishing activities and mean response of infauna and epifauna communities to fishing activities was much more negative in mud and sand communities than other habitats. Nephtys hombergii abundance also significantly decreased in areas of the Solent, UK, where bait digging (primarily for Nereis virens) had occurred (Watson et al., 2007). Similarly, Nephtys hombergii abundance was reduced by 50% in areas where tractor towed cockle harvesting was undertaken on experimental plots in Burry inlet, South Wales, and had not recovered after 86 days (Ferns et al., 2000).

    Sensitivity assessment. The biotopes will be impacted if damage to seabed surface features is widespread, however, motile and opportunistic fauna, such as Tubificoides benedii may recover quickly. Nephtys hombergii shows a greater negative impact (Collie et al., 2000; Ferns et al., 2000). Resistance is assessed as Low and resilience is assessed as High, providing a sensitivity assessment of Low.

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

    Activities that penetrate below the surface would remove a significant proportion of the characterizing species within the direct area of impact (see evidence under ‘abrasion’ above). Resistance of the biotopes is therefore assessed as Low, although the significance of the impact for the bed will depend on the spatial scale of the pressure footprint. Resilience is assessed as High, and sensitivity is assessed as Low.

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

    The biotopes are probably exposed to the high levels of suspended sediment characteristic of estuarine conditions. Therefore, the resident species are probably adapted to high suspended sediment levels. Aphelochaeta marioni lives infaunally and is a surface deposit feeder (Rayment, 2007a), therefore relying on a supply of nutrients at the sediment surface. An increased rate of siltation may result in an increase in food availability and therefore growth and reproduction of Aphelochaeta marioni. However, food availability would only increase if the additional suspended sediment contained a significant proportion of organic matter and the population would only be enhanced if food was previously limiting. A decrease in the suspended sediment would result in a decreased rate of deposition on the substratum surface and therefore a reduction in food availability for Aphelochaeta marioni. This would be likely to impair growth and reproduction.

    Tubificoides benedii displays preference for fine organic enriched sediments and may, therefore, increase in abundance if suspended solids settle and lead to increased organic enrichment (Gogina et al., 2010).

    Sensitivity assessment. Changes in light penetration or attenuation associated with this pressure are not relevant to Nephtys hombergii, Aphelochaeta marioni and Tubificoides benedii. As the species live in the sediment they are also likely to be adapted to increased suspended sediment (and turbidity). However, alterations in the availability of food or the energetic costs in obtaining food or changes in scour could either increase or decrease habitat suitability for these characterizing species. The following sensitivity assessment relies on expert judgement, utilizing evidence of species traits and distribution and therefore confidence has been assessed as low. Resistance is High as no significant negative effects are identified and potential benefits from increased food resources may occur. Resilience is also High as no recovery is required under the likely impacts. Sensitivity of the biotopes is therefore assessed as Not Sensitive.

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

    Aphelochaeta marioni lives infaunally in soft sediments and moves by burrowing. It deposit feeds at the surface by extending contractile palps from its burrow. An additional 5 cm layer of sediment would result in a temporary cessation of feeding activity, and therefore growth and reproduction are likely to be compromised. However, Aphelochaeta marioni would be expected to quickly relocate to its favoured depth, with no mortality. Kędra et al. (2010) reported Aphelochaeta mariori to occur in the Hornsundfjord, Svalbard, where sedimentation rates can vary between 0.1-35 cm/yr.

    Tubificoides live relatively deeply buried and can tolerate periods of low oxygen that may occur following the deposition of a fine layer of sediment. In addition, the presence of this species in areas experiencing deposition, such as estuaries, indicate that this species is likely to have a high tolerance to siltation events. Tubificoides spp. showed some recovery through vertical migration following the placement of a sediment overburden 6 cm thick on top of sediments (Bolam, 2011). Whomersley et al. (2010) experimentally buried plots on intertidal mudflats at two sites (Creeksea Crouch Estuary, England, and Blackness lower Forth Estuary, Scotland), where Tubificoides benedii were dominant species. For each treatment, anoxic mud was spread evenly to a depth of 4 cm on top of each treatment plot. The mud was taken from areas adjacent to the plots, and was obtained by scraping off the surface oxic layer and digging up the underlying mud from approximately 20 cm depth. Plots were subject to either low intensity treatments (burial every four weeks) or high (burial every two weeks). The experiment was carried out for 10 months at Creeksea and a year at Blackness. At Creeksea numbers of Tubificoides benedii increased in both burial treatments until the third month (high burial) and sixth month (low burial). At Blackness, increased numbers of Tubificoides benedii were found in both burial treatments after one month (Whomersley et al., 2010).

    In the low energy environment where the biotopes occur, a ‘light’ deposition of sediments is likely to be cleared in a few tidal cycles.

    Sensitivity assessment. The characterizing species Nephtys hombergii, Aphelochaeta marioni and Tubificoides benedii live in the sediment, to depths to 15 cm and are considered likely to resist smothering by 5 cm of sediment. Resistance and resilience are, therefore, assessed as High and the biotopes are considered Not Sensitive to a ‘light’ deposition of up to 5 cm of fine material in a single discrete event.

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

    Aphelochaeta marioni lives infaunally in soft sediments and moves by burrowing. It deposit feeds at the surface by extending contractile palps from its burrow. An additional layer of sediment would result in a temporary cessation of feeding activity, and therefore growth and reproduction are likely to be compromised. However, Aphelochaeta marioni would be expected to quickly relocate to its favoured depth, with no mortality. Kędra et al. (2010) reported Aphelochaeta mariori to occur in the Hornsundfjord, Svalbard, where sedimentation rates can vary between 0.1-35 cm/yr. Furthermore, Do et al. (2012) studied the macrobenthos recovery in the Arcachon Bay (France) following a deposition of sediment up to 10 cm thick that resulted from dredging activities. The authors reported Aphelochaeta marioni as considerably reduced or absent from impacted areas characterized mainly by mud substrata.

    The pressure benchmark (30 cm deposit) represents a significant burial event and the deposit may remain for some time in a sheltered mudflat. Some impacts on some characterizing species may occur and it is considered unlikely that significant numbers of the population could reposition, based on Bolam (2011). Polychaete species have been reported to migrate through depositions of sediment greater that the benchmark (30 cm of fine material added to the seabed in a single discrete event) (Maurer et al., 1982). However, it is not clear whether the characterizing species are likely to be able to migrate through a maximum thickness of fine sediment because muds tend to be more cohesive and compacted than sand. Some mortality of the characterizing species is likely to occur. Placement of the deposit will, therefore, result in a defaunated habitat until the deposit is recolonized.

    Sensitivity assessment. Resistance is assessed as Low and resilience as High and the biotopes are considered to have Low sensitivity to a ‘heavy’ deposition 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

    No Evidence is available on which to assess this pressure.

    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 is available on which to assess this pressure.

    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 species in the biotopes are likely to be sensitive to noise or vibration at the benchmark level and no information was found concerning the intolerance to noise. Species within the biotopes can probably detect vibrations caused by noise and in response may retreat in to the sediment for protection. However, the characterizing species are unlikely to be affected by noise pollution and so the biotopes are assessed as Not Sensitive.

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

    Changes in light availability impact primary production by phytoplankton in the water column and by the microphytobenthos on the sediment surface, which in turn may affect food availability for the characterizing species. Farke (1979) noted that Aphelochaeta marioni is intolerant to visual disturbance in a microsystem in the laboratory possibly due to its nocturnal life habits (Farke, 1979). In order to observe feeding and breeding in the microsystem, the animals had to be gradually acclimated to lamp light. Even then, additional disturbance, such as an electronic flash, caused the retraction of palps and cirri and cessation of all activity for some minutes. Visual disturbance, in the form of direct illumination during the species' active period at night, may therefore result in loss of feeding opportunities, which may compromise growth and reproduction.

    Sensitivity assessment. However, the characterizing community of the biotopes live infaunally and are unlikely to be directly dependent on sunlight so the biotopes are considered to have High resistance and, by default, High resilience and therefore is Not Sensitive to the introduction of light or shading.

    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.

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

    Most species will respond to the shading caused by the approach of a predator, however, their visual acuity is probably very low. Farke (1979) noted that Aphelochaeta marioni is intolerant to visual disturbance in a microsystem in the laboratory possibly due to its nocturnal life habits (Farke, 1979). In order to observe feeding and breeding in the microsystem, the animals had to be gradually acclimated to lamp light. Even then, additional disturbance, such as an electronic flash, caused the retraction of palps and cirri and cessation of all activity for some minutes.

    Most aquatic oligochaetes, such as Tubificoides spp. have no eyes, although a few have simple ocelli (eyspots) which are light receptors. However, as the characterizing species live in the sediment, the species will most probably not be impacted at the pressure benchmark.

    Sensitivity assessment.  At the pressure benchmark, resistance is assessed as High and resilience as High and the biotopes are assessed as Not Sensitive.

    Biological Pressures

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

    The important characterizing species in the biotopes are not cultivated or likely to be translocated. This pressure is therefore considered Not Relevant.

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

    The biotopes may be exposed to a number of invasive species that can cause impacts. The largest effect may be from species that significantly alter the character of the biotopes, such as reef-forming species and invasive vegetation. Commito (1987) found that the population density of Tubificoides benedii was the same or higher in mussel beds than in open areas, suggesting that colonization of sediments by the pacific oyster Crassostrea gigas would not necessarily impact the population (although it would alter the character of the biotopes). Tang & Kristensen (2010) found that abundance of macrofauna, including Tubificoides, was lower in marsh invaded by the hybrid cordgrass Spartina anglica than in mudflats. Colonization of upper mudflats by this species would alter the character of the biotopes resulting in loss and reclassification.

    Sensitivity assessment. The biotopes may be sensitive to invasion by Spartina anglica which would alter the character of the mudflat and the biological assemblage. Resistance is assessed as None and resilience as Very Low as the biotopes will not recover unless the INIS is removed. Sensitivity is therefore assessed as High.

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

    Introduced organisms (especially parasites or pathogens) are a potential threat in all coastal ecosystems. Little information was found regarding microbial infection of polychaetes, although Gibbs (1971) recorded that nearly all of the population of Aphelochaeta marioni in Stonehouse Pool, Plymouth Sound, was infected with a sporozoan parasite belonging to the acephaline gregarine genus Gonospora, which inhabits the coelom of the host. No evidence was found to suggest that gametogenesis was affected by Gonospora infection and there was no apparent reduction in fecundity.

    Marine oligochaetes host numerous protozoan parasites without apparent pathogenic effects even at high infestation levels (Giere & Pfannkuche, 1982).

    Sensitivity assessment. The biotopes are judged to have High resistance to this pressure. By default, resilience is assessed as High and the biotopes are classed as Not Sensitive.

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

    Nephtys hombergii is directly removed through commercial bait digging and by recreational anglers and abundance significantly decreased in areas of the Solent, UK, where bait digging (primarily for Nereis virens) had occurred (Watson et al., 2007).

    Recovery of Nephtys hombergii has been assessed to be very high as repopulation would initially occur relatively rapidly via adult migration and later by larval recruitment. Dittman et al. (1999) observed that Nephtys hombergii was amongst the macrofauna that colonized experimentally disturbed tidal flats within two weeks of the disturbance that caused defaunation of the sediment. However, if sediment is damaged recovery is likely to be slower, for instance Nephtys hombergii abundance was reduced by 50% in areas where tractor towed cockle harvesting was undertaken on experimental plots in Burry inlet, South Wales, and had not recovered after 86 days (Ferns et al., 2000).

    Sensitivity assessment. Resistance is Low for SS.SMu.SMuVS.NhomTubi due to direct removal of a characterizing species Nephtys hombergii, that on commercial scales can remove a large proportion of the population. For instance, Neves de Carvalho et al. (2013) calculated that bait digging activities in the Douro estuary, Portugal may remove up to 9.9 tonnes of Hediste diversicolor. Resilience is assessed as Medium, as regions that are not regularly harvested may recover rapidly but continued harvesting will impact the population. Sensitivity of SS.SMu.SMuVS.NhomTubi is assessed as Medium. It is important to consider that the spatial extent and duration of harvesting is important to consider when assessing this pressure, as smaller scale extraction may not impact the entire extent of the biotope but greater scale extraction over a long period would cause longer term impacts.

    On the other hand, it is extremely unlikely that species indicative of sensitivity of SS.SMu.SMuVS.AphTubi would be targeted for extraction, so this pressure is therefore considered Not Relevant for SS.SMu.SMuVS.AphTubi.

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

    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. Species in the biotopes, including the characterizing species, may be damaged or directly removed by static or mobile gears that are targeting other species (see abrasion and penetration pressures). Hall & Harding (1997) demonstrated that commercial cockle harvesting by suction dredging had significant effects on soft-sediment infaunal communities. Following dredging, species numbers were reduced by up to 30% and abundances by up to 50%.

    Collie et al. (2000) identified that well established intertidal communities suffered the greatest impact from bottom towed fishing activities. The review concluded that there were ecologically important impacts from removal of >50% of fauna from bottom towed fishing activity (dredge and trawls) (Collie et al., 2000). However, the burrowing and tunnelling traits of the characterizing species may provide some resistance to this pressure. Kaiser et al. (2001) carried out experimental hand raking, similar to that used in intertidal cockle fisheries. Both small and large raked plots showed changed communities in comparison to control plots, smaller plots recovered in 56 days, whilst larger plots remained in an altered state.

    Collie et al. (2000) found that Nephtys hombergii abundance was negatively affected by fishing activities. Mean response of infauna and epifauna communities to fishing activities was also much more negative in mud and sand communities than other habitats. Nephtys hombergii abundance also significantly decreased in areas of the Solent, UK, where bait digging (primarily for Nereis virens) had occurred (Watson et al., 2007). Similarly, Nephtys hombergii abundance was reduced by 50% in areas where tractor towed cockle harvesting was undertaken on experimental plots in Burry inlet, South Wales, and had not recovered after 86 days (Ferns et al., 2000).

    Aphelochaeta marioni is a soft bodied organism which exposes its palps and cirri at the surface while feeding (Rayment, 2007a). The species lives infaunally in soft sediment, usually within a few centimetres of the sediment surface. Physical disturbance, such as dredging or dragging an anchor, would be likely to penetrate the upper few centimetres of the sediment and cause physical damage to Aphelochaeta marioni.

    Sensitivity assessment. Resistance is Low due to direct removal or damage of characterizing species that on commercial scales can remove a large proportion of the population. Resilience is assessed as Medium as regions that are not regularly harvested may recover rapidly but continued harvesting will impact the population. Sensitivity is assessed as Medium. It is important to consider that the spatial extent and duration of areas impacted by fishing gear effort is important to consider when assessing this pressure as smaller scale extraction may not impact the entire extent of the biotopes but greater scale extraction over a long period would cause longer term impacts. The type of fishing activity is also important to consider in relation to the type and severity of the impact.

    Importance review

    Policy/Legislation

    Habitats Directive Annex 1Estuaries

    Exploitation

    None known.

    Additional information

    No additional information.

    Bibliography

    1. Alheit, J., 1978. Distribution of the polychaete genus Nephtys: a stratified random sampling survey. Kieler Meeresforschungen, 4, 61-67.
    2. Arndt, C. & Schiedek, D., 1997. Nephtys hombergii, a free living predator in marine sediments: energy production under environmental stress. Marine Biology, 129, 643-540.
    3. Bagheri, E. & McLusky, D., 1982. Population dynamics of oligochaetes and small polychaetes in the polluted forth estury ecosystem. Netherlands Journal of Sea Research, 16, 55-66.

    4. 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.
    5. Barnes, R.S.K. & Hughes, R.N., 1992. An introduction to marine ecology. Oxford: Blackwell Scientific Publications.
    6. Barnes, R.S.K., 1994. The brackish-water fauna of northwestern Europe. Cambridge: Cambridge University Press.
    7. 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.
    8. Bentley, M.G., Olive, P.J.W., Garwood, P.R. & Wright, N.H., 1984. The spawning and spawning mechanism of Nephtys caeca (Fabricius, 1780) and Nephtys hombergii Savigny, 1818 (Annelida: Polychaeta). Sarsia, 69, 63-68.
    9. Beukema, J.J., 2002. Expected changes in the benthic fauna of Wadden Sea tidal flats as a result of sea-level rise or bottom subsidence. Journal of Sea Research, 47 (1), 25-39.
    10. 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.

    11. 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.
    12. Birtwell, I.K. & Arthur, D.R., 1980. The ecology of tubificids in the Thames Estuary with particular reference to Tubifex costatus (Claparède). In Proceedings of the first international symposium on aquatic oligochaete biology, Sydney, British Colombia, Canada, May 1-4, 1979. Aquatic oligochaete biology (ed. R.O. Brinkhurst & D.G. Cook), pp. 331-382. New York: Plenum Press
    13. Bolam, S. & Whomersley, P., 2003. Invertebrate recolonization of fine-grained beneficial use schemes: An example from the southeast coast of England. Journal of Coastal Conservation, 9 (2), 159-169.

    14. Bolam, S.G., 2011. Burial survival of benthic macrofauna following deposition of simulated dredged material. Environmental Monitoring and Assessment, 181 (1-4), 13-27.
    15. 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.
    16. 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.
    17. 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.
    18. 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.]
    19. 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.
    20. 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.
    21. 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.
    22. Cardell, M.J., Sarda, R. & Romero, J., 1998. Spatial changes in sublittoral soft-bottom polychaete assemblages due to river inputs and sewage discharges. Acta Oecologica, 20, 343-351.
    23. Clark, R.B. & Haderlie, E.C., 1960. The distribution of Nephtys cirrosa and Nephtys hombergii of the south western coasts of England and Wales. Journal of Animal Ecology, 29, 117-147.
    24. Clark, R.B., Alder, R.R. & McIntyre, A.D., 1962. The distribution of Nephtys sp. on the Scottish coast. Journal of Animal Ecology, 31, 359-372.
    25. Coates, D.A., Deschutter, Y., Vincx, M. & Vanaverbeke, J., 2014. Enrichment and shifts in macrobenthic assemblages in an offshore wind farm area in the Belgian part of the North Sea. Marine Environmental Research, 95, 1-12.
    26. 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.
    27. 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.
    28. Commito, J.A. & Boncavage, E.M., 1989. Suspension-feeders and coexisting infauna: an enhancement counterexample. Journal of Experimental Marine Biology and Ecology, 125 (1), 33-42.

    29. Commito, J.A., 1987. Adult-larval interactions: predictions, mussels and cocoons. Estuarine, Coastal and Shelf Science, 25, 599-606.
    30. Compton, T.J., Holthuijsen, S., Koolhaas, A., Dekinga, A., Ten Horn, J., Smith, J., Galama, Y., Brugge, M., van der Wal, D., Van der Meer, J., Van Der Veer, H.W. & Piersma, T., 2013. Distinctly variable mudscapes: Distribution gradients of intertidal macrofauna across the Dutch Wadden Sea. Journal of Sea Research, 82, 103-116.

    31. 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. Joint Nature Conservation Committee, Peterborough. www.jncc.gov.uk/MarineHabitatClassification.
    32. 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.
    33. Covazzi-Harriague, A., Misic, C., Petrillo, M. & Albertelli, G., 2007. Stressors affecting the macrobenthic community in Rapallo harbour (Ligurian Sea, Italy). Scientia Marina, 71 (4), 705-714.
    34. 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.
    35. Daro, M.H. & Polk, P., 1973. The autecology of Polydora ciliata along the Belgian coast. Netherlands Journal of Sea Research, 6, 130-140.
    36. 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.
    37. 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.
    38. 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.
    39. De Biasi, A. & Pacciardi, L., 2008. Macrobenthic communities in a fishery exclusion zone and in a trawled area of the middle Adriatic Sea (Italy). Ciencias Marinas, 34 (4).
    40. Degraer, S., Wittoeck, J., Appeltans, W., Cooreman, K., Deprez, T., Hillewaert, H., Hostens, K., Mees, J., Berge, V. & Vincx, M., 2006. The macrobenthos atlas of the Belgian part of the North Sea. Belgian Science Policy.
    41. Dernie, K.M., Kaiser, M.J., Richardson, E.A. & Warwick, R.M., 2003. Recovery of soft sediment communities and habitats following physical disturbance. Journal of Experimental Marine Biology and Ecology, 285-286, 415-434.
    42. 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.
    43. 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.
    44. 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.
    45. Do, V.T., de Montaudouin, X., Blanchet, H. & Lavesque, N., 2012. Seagrass burial by dredged sediments: Benthic community alteration, secondary production loss, biotic index reaction and recovery possibility. Marine Pollution Bulletin, 64 (11), 2340-2350.
    46. Emery, K.O. & Stevenson, R.E., 1957. Estuaries and lagoons. In Treatise on marine ecology and paleoecology.1. Ecology, (ed. J.W. Hedgpeth), USA: Geological Society of America.
    47. Fallesen, G. & Jørgensen, H.M., 1991. Distribution of Nephtys hombergii and Nephtys ciliata (Polychaeta: Nephtyidae) in Århus Bay, Denmark, with emphasis on the severe oxygen deficiency. Ophelia, Supplement 5, 443-450.
    48. 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.
    49. 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.
    50. Fish, J.D. & Fish, S., 1996. A student's guide to the seashore. Cambridge: Cambridge University Press.
    51. 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.
    52. 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.
    53. Gibbs, P.E., Langston, W.J., Burt, G.R. & Pascoe, P.L., 1983. Tharyx marioni (Polychaeta) : a remarkable accumulator of arsenic. Journal of the Marine Biological Association of the United Kingdom, 63, 313-325.
    54. Giere, O., 1977. An ecophysiological approach to the microdistribution of meiobenthic Oligochaeta. I. Phallodrilus monospermathecus (Knöllner)(Tubificidae) from a subtropical beach at Bermuda. Biology of benthic organisms. Pergamon Press New York, 285-296.

    55. Giere, O., 1992. Benthic life in sulfidic zones of the sea-ecological and structural adaptations to a toxic environment. Verhandlungen der Deutschen Zoologischen Gesellschaft, 85 (2), 77-93.
    56. Giere, O., 2006. Ecology and biology of marine oligochaeta–an inventory rather than another review. Hydrobiologia, 564 (1), 103-116.

    57. Giere, O. & Pfannkuche, O., 1982. Biology and ecology of marine Oligochaeta, a review. Oceanography and Marine Biology, 20, 173-309.

    58. 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.
    59. Gillett, D.J., Holland, A.F. & Sanger, D.M., 2007. On the ecology of oligochaetes: monthly variation of community composition and environmental characteristics in two South Carolina tidal creeks. Estuaries and Coasts, 30 (2), 238-252.

    60. 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. 
    61. Gray, J.S., 1971. The effects of pollution on sand meiofauna communities. Thalassia Jugoslovica, 7, 76-86.
    62. 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.
    63. 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.
    64. Hall, J.A. & Frid, C.L.J. 1998. Colonisation patterns of adult macrobenthos in a polluted North Sea Estuary. Aquatic Ecology, 31, 333-340.
    65. 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.
    66. Hall, S.J. & Harding, M.J.C., 1997. Physical disturbance and marine benthic communities: the effects of mechanical harvesting of cockles on non-target benthic infauna. Journal of Applied Ecology, 34, 497-517.
    67. Haskoning UK Ltd. 2006. Investigation into the impact of marine fish farm deposition on maerl beds. Scottish Natural Heritage Commissioned Report No. 213 (ROAME No. AHLA10020348).

    68. 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.
    69. 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.
    70. Hunter, J., & Arthur, D.R., 1978. Some aspects of the ecology of Peloscolex benedeni Udekem (Oligochaeta: Tubificidae) in the Thames estuary. Estuarine and Coastal Marine Science, 6, 197-208.
    71. Huthnance, J., 2010. Ocean Processes Feeder Report. London, DEFRA on behalf of the United Kingdom Marine Monitoring and Assessment Strategy (UKMMAS) Community.
    72. 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.
    73. 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,
    74. Kaiser, M., Broad, G. & Hall, S., 2001. Disturbance of intertidal soft-sediment benthic communities by cockle hand raking. Journal of Sea Research, 45 (2), 119-130.
    75. Kędra, M., Gromisz, S., Jaskuła, R., Legeżyńska, J., Maciejewska, B., Malec, E., Opanowski, A., Ostrowska, K., Włodarska-Kowalczuk, M. & Węsławski, J., 2010. Soft bottom macrofauna of an All Taxa Biodiversity Site: Hornsund (77○ N, Svalbard). Polish Polar Research, 31 (4), 309-326.
    76. 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
    77. 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.
    78. Linke, O., 1939. Die Biota des Jadebusenwatts. Helgolander Wissenschaftliche Meeresuntersuchungen, 1, 201-348.
    79. 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
    80. Markert, A., Wehrmann, A. & Kröncke, I., 2010. Recently established Crassostrea-reefs versus native Mytilus-beds: differences in ecosystem engineering affects the macrofaunal communities (Wadden Sea of Lower Saxony, southern German Bight). Biological Invasions, 12 (1), 15-32.
    81. Mathivat-Lallier, M.H. & Cazaux, C., 1991. Life-history of Nephtys hombergii in Arcachon Bay. Estuarine and Coastal Marine Science, 32, 1-9.
    82. 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.

    83. 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.
    84. 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.
    85. McLusky, D.S., Teare, M. & Phizachlea, P., 1980. Effects of domestic and industrial pollution on distribution and abundance of aquatic oligochaetes in the Forth estuary. Helgolander Wissenschaftliche Meeresuntersuchungen, 33, 384-392.
    86. 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.
    87. Murina, V., 1997. Pelagic larvae of Black Sea Polychaeta. Bulletin of Marine Science, 60, 427-432.
    88. 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.
    89. Nilsson, H.C. & Rosenberg, R., 1994. Hypoxic response of two marine benthic communities. Marine Ecology Progress Series, 115, 209-217.
    90. 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.
    91. 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.
    92. 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.
    93. 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.
    94. Orvain, F., Sauriau, P.-G., Le Hir, P., Guillou, G., Cann, P. & Paillard, M., 2007. Spatio-temporal variations in intertidal mudflat erodability: Marennes-Oléron Bay, western France. Continental Shelf Research, 27 (8), 1153-1173.
    95. Oyenekan, J.A., 1986. Population dynamics and secondary production of Nephtys hombergii (Polychaeta: Nephtyidae). Marine Biology, 93, 217-223.
    96. 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.
    97. 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.
    98. 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.
    99. Rayment, W.J. 2007a. Aphelochaeta marioni A bristleworm. 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/species/detail/1556

    100. 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.
    101. 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.
    102. Rodrigues, A.M., Quintino, V., Sampaio, L., Freitas, R. & Neves, R., 2011. Benthic biodiversity patterns in Ria de Aveiro, Western Portugal: Environmental-biological relationships. Estuarine, Coastal and Shelf Science, 95 (2–3), 338-348.
    103. Rygg, B., 1985. Effect of sediment copper on benthic fauna. Marine Ecology Progress Series, 25, 83-89.
    104. Schöttler, U., 1982. An investigation on the anaerobic metabolism of Nephtys hombergii (Annelida: Polychaeta). Marine Biology, 71 (3), 265-269.
    105. Schueckel, U. & Kroencke, I., 2013. Temporal changes in intertidal macrofauna communities over eight decades: A result of eutrophication and climate change. Estuarine Coastal and Shelf Science, 117, 210-218.
    106. Sheehan, E., Coleman, R., Thompson, R. & Attrill, M., 2010. Crab-tiling reduces the diversity of estuarine infauna. Marine Ecology Progress Series, 411, 137-148.

    107. Suchanek, T.H., 1993. Oil impacts on marine invertebrate populations and communities. American Zoologist, 33, 510-523.
    108. Sundborg, Å., 1956. The River Klarälven: a study of fluvial processes. Geografiska Annaler, 38 (2), 125-237.

    109. Tang, M. & Kristensen, E., 2010. Associations between macrobenthos and invasive cordgrass, Spartina anglica, in the Danish Wadden Sea. Helgoland Marine Research, 64 (4), 321-329.

    110. Thiermann, F., Niemeyer, A-S. & Giere, O., 1996. Variations in the sulfide regime and the distribution of macrofauna in an intertidal flat in the North Sea. Helgolander Meeresuntersuchungen, 50, 87-104.
    111. UKTAG, 2014. UK Technical Advisory Group on the Water Framework Directive [online]. Available from: http://www.wfduk.org
    112. Vader, W.J.M., 1964. A preliminary investigation in to the reactions of the infauna of the tidal flats to tidal fluctuations in water level. Netherlands Journal of Sea Research, 2, 189-222.
    113. Van Colen, C., De Backer, A., Meulepas, G., van der Wal, D., Vincx, M., Degraer, S. & Ysebaert, T., 2010a. Diversity, trait displacements and shifts in assemblage structure of tidal flat deposit feeders along a gradient of hydrodynamic stress. Marine Ecology Progress Series, 406, 79-89.

    114. Van Hoey, G., Guilini, K., Rabaut, M., Vincx, M. & Degraer, S., 2008. Ecological implications of the presence of the tube-building polychaete Lanice conchilega on soft-bottom benthic ecosystems. Marine Biology, 154 (6), 1009-1019.

    115. Verdonschot, P., Smies, M. & Sepers, A., 1982. The distribution of aquatic oligochaetes in brackish inland waters in the SW Netherlands. Hydrobiologia, 89 (1), 29-38.

    116. Vismann, B., 1990. Sulphide detoxification and tolerance in Nereis (Hediste) diversicolor and Nereis (Neanthes) virens (Annelida: Polychaeta). Marine Ecology Progress Series, 59, 229-238.
    117. Waldock, R., Rees, H.L., Matthiessen, P. & Pendle, M.A., 1999. Surveys of the benthic infauna of the Crouch Estuary (UK) in relation to TBT contamination. Journal of the Marine Biological Association of the United Kingdom, 79, 225 - 232.
    118. 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.
    119. Whomersley, P., Huxham, M., Bolam, S., Schratzberger, M., Augley, J. & Ridland, D., 2010. Response of intertidal macrofauna to multiple disturbance types and intensities – an experimental approach. Marine Environmental Research, 69 (5), 297-308.

    120. Wilson, D.P., 1936. Notes on the early stages of two polychaetes, Nephtys hombergii Lamarck and Pectinaria koreni Malmgren. Journal of the Marine Biological Association UK, 21, 305-310.
    121. Wolff, W.J., 1971b. Changes in intertidal benthos communities after an increase in salinity. Thalassia Jugoslavica, 7, 429-434.
    122. 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:

    De-Bastos, E. & Hiscock, K. 2016. 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: 29/06/2016