|Researched by||Georgina Budd||Refereed by||Mike Kendall|
|Authority||(O.F. Müller, 1776)|
|Other common names||-||Synonyms||Nereis diversicolor (O.F. Müller, 1776), Nereis (Hediste) diversicolor (O.F. Müller, 1776)|
Hediste diversicolor is one of the commonest intertidal polychaetes in estuaries. Its body appears flattened with a prominent dorsal blood vessel. Adults may reach 6-12 cm in length and consist of between 90-120 chaetae bearing segments (chaetigers). Appendages on the head are conspicuous consisting of two antennae, and two palps, and four pairs of tentacles. The paired parapodia have dorsal and ventral chaetae and are used for crawling and swimming. The colour of Hediste diversicolor varies. Mature worms become a brighter green approaching and during spawning, otherwise specimens appear to be a reddish orange or brown. There has been considerable controversy over the name to be applied to this species and Nereis diversicolor is used by many authors.
The form and distribution of paragnaths on the pharynx can be very useful in identification and Kinberg (1866, cited in Chambers & Garwood, 1992) assigned roman numerals to eight different areas of the pharynx that bear paragnaths. However, the number of paragnaths can vary considerably both within and between populations and this variation is thought to be a result of habitat and feeding preferences (Barnes & Head, 1977).
See Chambers & Garwood (1992) for further description and detail on identification.
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|Phylum||Annelida||Segmented worms e.g. ragworms, tubeworms & fanworms|
|Class||Polychaeta||Bristleworms, e.g. ragworms, scaleworms, paddleworms, fanworms and tubeworms|
|Authority||(O.F. Müller, 1776)|
|Recent Synonyms||Nereis diversicolor (O.F. Müller, 1776)Nereis (Hediste) diversicolor (O.F. Müller, 1776)|
|Typical abundance||High density|
|Male size range||60-120mm|
|Male size at maturity||60-70mm|
|Female size range||60-70mm|
|Female size at maturity|
|Growth form||Vermiform segmented|
|Body flexibility||High (greater than 45 degrees)|
|Characteristic feeding method||Non-feeding, Passive suspension feeder, Scavenger, Sub-surface deposit feeder, Surface deposit feeder|
|Typically feeds on||Mud, sand & detritus. Phytoplankton & plankton. Other macrofauna.|
|Is the species harmful?||No|
Hediste diversicolor is omnivorous and exhibits a diversity of feeding modes; carnivory, scavenging, filter feeding on suspended particles and deposit-feeding on materials in and on the surface layers of the sediment (Barnes, 1994). Hediste diversicolor feeds using an eversible pharynx and the sensory appendages on the head, namely palps and tentacles (M. Kendall, pers. comm.). < A conspicuous difference between Hediste diversicolor and the closely related polychaete Nereis virens is the unique ability of Hediste diversicolor to satisfy its metabolic requirements from a diet of phytoplankton, like a typical obligate filter-feeder (Nielsen et al., 1995).
The filter feeding mechanism was described by Harley (1950). A funnel-shaped net consisting of fine mucous threads is drawn across the burrow and a water current is driven through the net by undulating body movements (Fauchald & Jumars, 1979). This is best observed in a tank (M. Kendall, pers. comm.). When sufficient particles have accumulated on the net, they are consumed along with the entire net (Fauchald & Jumars, 1979). After an interval, the net is replaced (M. Kendall, pers. comm.). Riisgård (1991) suspected that Hediste diversicolor is a hitherto undervalued key organism in the control of phytoplankton in shallow brackish waters. It is unknown to what extent Hediste diversicolor utilizes its potential to subsist on suspended food particles in nature but can be considered a suspension feeder when a sufficient number of algal cells are present in the water (Riisgård, 1991).
When deposit feeding, Esnault et al. (1990) recognized two main types of searching behaviour exhibited by Hediste diversicolor. The first involved the worm crawling on the surface of the substratum prospecting for food, catching it with its jaws and ingesting it immediately. The second type saw the worm depositing a string of mucous on either side of its body on the substrate surface. When the worm retreated back into its burrow the mucous was brought back and built it into a pellet which can be consumed there and then or stored for consumption later on (Esnault et al., 1990).
Olivier et al. (1995) found that juvenile Hediste diversicolor can select detritus on the sediment surface and accumulate it in their burrow. The juveniles irrigate the burrows thereby maintaining an aerobic condition that favours the decaying process of the plant debris by stimulating bacterial growth ('gardening').
Lucas & Bertru (1997) found bacteriolytic activity in the digestive system of Hediste diversicolor thus highlighting the ability of this species to feed on bacteria.
The variable colours of Hediste diversicolor approaching maturity and during spawning (see reproduction) are due to varying proportions of green (biliverdin), orange and brown (carotenoids) pigments. The green colour of mature males and females is caused by biliverdin present in the gut wall, the epidermis and coelomic cells and is formed by the breakdown of haemoglobin in the blood. In males, the white mass of sperm in the coelom gives it a lighter green colour (Dales, 1950). In mature specimens during and after spawning, the green appearance is also enhanced by a complete extraction of carotenoids from the body wall (Dales & Kennedy, 1954).
|Physiographic preferences||Ria / Voe, Estuary, Enclosed coast / Embayment|
|Biological zone preferences||Lower eulittoral, Lower littoral fringe, Mid eulittoral, Upper eulittoral, Upper littoral fringe|
|Substratum / habitat preferences||Mud, Muddy sand, Sandy mud|
|Tidal strength preferences||Very Weak (negligible), Weak < 1 knot (<0.5 m/sec.)|
|Wave exposure preferences||Extremely sheltered, Sheltered, Very sheltered|
|Salinity preferences||Low (<18 psu), Reduced (18-30 psu), Variable (18-40 psu)|
|Other preferences||No text entered|
|Migration Pattern||Non-migratory / resident|
Distribution & density
Hediste diversicolor is an euryhaline species and can withstand great variances in salinity. Smith (1956) reported that, in the Tamar estuary, England, individuals of this species living at the upstream limit regularly experience salinities less than 0.5 ppt. In marine dominated habitats, Hediste diversicolor behaves as a brackish water animal and is found in the least saline portion of the available ground (Smith, 1956). The distribution of Hediste diversicolor in high salinity areas is likely to be reduced as result of competition in the form of interspecific aggressions (Kristensen, 1988). In a study focussing on the distribution of nereid polychaetes in Danish coastal waters, Kristensen (1988) found that Hediste diversicolor could only maintain high population densities in marginal environments when the fitness of stronger competitors such as Nereis virens was reduced.
In estuaries the maximum density of the Hediste diversicolor population normally occurs in the middle regions, with density decreasing both towards the head and mouth of the estuary. Smith (1956), found that the maximum population density of Hediste diversicolor in the Tamar estuary corresponded to that portion of the estuary with the greatest salinity variation. The density of worms varies between locations and throughout the reproductive cycle. Numbers of juveniles may be over 100 000 per m² (Clay, 1967(c)). In the Ythan Estuary, Scotland, the density of adult Hediste diversicolor was reported to be 961 per m² (Chambers & Milne, 1975).Burrows
|Reproductive type||Gonochoristic (dioecious)|
|Reproductive frequency||Semelparous / monotely|
|Fecundity (number of eggs)||1,000-10,000|
|Generation time||1-2 years|
|Age at maturity||See additional text|
|Season||Spring - See additional text|
|Life span||See additional information|
|Duration of larval stage||Not relevant|
|Larval dispersal potential||0 - 10 km|
|Larval settlement period||Not relevant|
This MarLIN sensitivity assessment has been superseded by the MarESA approach to sensitivity assessment. MarLIN assessments used an approach that has now been modified to reflect the most recent conservation imperatives and terminology and are due to be updated by 2016/17.
|Hediste diversicolor is infaunal and is reliant upon a muddy / sandy sediment in which to burrow. Physical removal of the substratum e.g. as a result of channel dredging activities would remove with it the entire associated population of Hediste diversicolor. The ability of postlarvae and larger juveniles and adults of Hediste diversicolor to swim, burrow and be carried by bedload transport can aid the rapid recolonization of disturbed sediments (Shull, 1997). Davey & George (1986), found evidence that larvae of Hediste diversicolor were tidally dispersed within the Tamar Estuary over a distance of 3 km and well away from areas of dense adult populations. However, this dispersal may not always lead the larvae to a favourable area and it is likely that there will be some loss.|
|Hediste diversicolor inhabits depositional environments. It is capable of burrowing to depths of up to 0.3 m and reworking sub-surface modifications of its burrow through fine clays and sand. Smith (1955) found no appreciable difference in the population of a Hediste diversicolor colony which had been covered by several inches of sand through which the worms tunneled. It would not be adversely affected by smothering with additional sediments. However, smothering with impermeable materials would prevent Hediste diversicolor clearing the burrow to the sediment surface and prevent feeding. Larvae are more intolerant than adults as they are still acquiring the physical ability to burrow (see larval sensitivity).|
|Tolerant*||Not relevant||Not sensitive*||Low|
|Increased siltation maybe beneficial to feeding. As a surface-deposit feeder and suspension feeder Hediste diversicolor will be able to utilize suspended matter as a food resource. Increased deposition of silt onto the mudflats can raise the height of the mudflats and therefore increasing the exposure time of infaunal communities at low tide (Jones et al., 2000). At the benchmark level however, this is unlikely to have an adverse effect on Hediste diversicolor.|
|Hediste diversicolor inhabits a burrow within the sediment which may be up to 0.3m deep. The species retreats within the burrow during periods of exposure and thus away from the desiccating factors of sunlight and wind. Residual surface and interstitial water prevent the burrow and thus Hediste diversicolor from drying. Therefore Hediste diversicolor is largely able to avoid desiccation. Specimens found at the upper limits in the intertidal zone may become stressed by desiccation if the substratum begins to dry, but Hediste diversicolor is sufficiently mobile to retreat back to damper substrata . Consequently this species is considered to have a low intolerance to the benchmark change in desiccating factors.|
|Hediste diversicolor inhabits a burrow within the sediment which may be up to 0.3m deep. The species retreats within the burrow during periods of exposure and thus away from desiccating factors of sunlight and wind. Thus Hediste diversicolor can avoid some detrimental aspects of emergence (see desiccation). However, whilst in retreat within the burrow during extended periods of emergence, the ragworm is prevented from actively feeding at the surface and is likely to be prone to more intense predation pressure from wading birds as they have longer to search the mudflats. An increased emergence regime is also likely to cause a decline in the abundance of ragworms at the upper limits of the intertidal zone, as they may become stressed by desiccation if the substrata begin to dry and are prone to more extremes of temperature, but Hediste diversicolor is sufficiently mobile to gradually retreat back to damper substrata. Consequently, this species is considered to have a low intolerance to the benchmark change in emergence.|
|Hediste diversicolor characteristically inhabits littoral mudflats predominantly of clay (particles < 4 µm), silt (4-63 µm) and to a lesser extent very fine sand (63-125 µm) (Jones et al., 2000). The type direction and speed of the currents control sediment deposition within an area. A change in two categories in water flow rate from weak and negligible to moderately strong and strong would entrain and maintain particles in suspension and erode the mud. As a result the scouring and consequent redistribution of components of the substratum would alter the extent of suitable habitat available to populations of Hediste diversicolor. Recovery of this species would be influenced by the length of time it would take for the potential habitat to return to a suitable state for recolonization by adult and juvenile specimens from adjacent habitats, and the establishment of a breeding population. This may take between one and three years, as populations differ in reaching maturity (Dales, 1950; Mettam et al., 1982; Olive & Garwood, 1981), from the time that the habitat again becomes suited to the species.|
|The geographic range of Hediste diversicolor (see adult distribution) suggests that it is tolerant of a range of temperatures and a long term chronic temperature increase or decrease is unlikely to have an adverse effect on UK populations. Hediste diversicolor can tolerate temperatures from below zero under Baltic ice to high summer temperatures in Black Sea lagoons (Smith, 1977).|
A decrease in temperature has been shown to be beneficial to Hediste diversicolor through reduction in numbers of their predators. A severe winter in the Wadden Sea in 1995/1996 saw an increased abundance of this species coinciding with a reduction in the numbers of Carcinus maenus and Crangon crangon (Armonies et al., 2001). A similar increase in abundance was noted in the same area between 1978 and 1987 after a series of cold winters: mean density increased from 24 / m² to 151 / m² respectively (Beukema, 1990).
Species dwelling in the sediments are likely to be protected from direct effects of temperature change at the surface, for instance Hediste diversicolor burrows deeper in very cold and frosty weather (Linke, 1939). In addition, insensitivity to temperature change is limited by the insulating properties of the mud in which it lives (M. Kendall, pers. comm.).
Temperature change may adversely affect reproduction. Bartels-Hardege & Zeeck (1990) demonstrated that an increase from 12°C and maintenance of water temperature at 16°C induced reproduction in specimens outside the normal period of spawning (see reproduction), and without a drop in temperature to simulate winter conditions the spawning period was prolonged and release of gametes was not synchronized. Poor synchronization of spawning could result in reduced recruitment, as gametes are wasted and mature specimens die shortly after gamete release. Therefore, an intolerance of intermediate has been recorded.
|Hediste diversicolor characteristically inhabits estuaries where turbidity is typically higher than other coastal waters. Changes in the turbidity may influence the abundance of phytoplankton available as a food source that may be attained through filter feeding. However, Hediste diversicolor utilizes various other feeding mechanisms and, at the benchmark level, the likely effects of a change in turbidity are limited.|
|Hediste diversicolor is infaunal, inhabiting a burrow in which it seeks refuge from predators and may partially emerge to seek and capture food. In addition, it inhabits low energy depositional environments. An alteration of factors within the environment that increases wave exposure is likely to cause erosion of the substrata and consequently, loss of habitat. |
Recovery would be influenced by the length of time it would take for the habitat to return to a suitable state for recolonization by adult and juvenile specimens from adjacent habitats, and the establishment of a breeding population. This may take between one and three years, as populations differ in reaching maturity (Dales, 1950; Mettam et al., 1982; Olive & Garwood, 1981), from the time that the habitat again becomes suited to the species.
|Tolerant*||Not relevant||Not sensitive*||Low|
|Hediste diversicolor may be able to detect some noise vibration but is not known to exhibit a significant response at the benchmark levels. However, wildfowl which prey upon estuarine infauna such as Hediste diversicolor are known to be disturbed by noise, consequently predation pressure upon Hediste diversicolor may be reduced for the length of time that the disturbance continues.|
|Tolerant||Not relevant||Not sensitive||High|
|Hediste diversicolor demonstrates a distinct movement towards darkness (skototaxis) and it has been shown that feeding sensitizes the worm to light and influences their response to a sudden increase in illumination (Herter, 1926; Evans, 1966, in Clay 1967(c)). Otherwise, Hediste diversicolor lacks the visual ability to be affected by the visual presence of moving objects not normally found in the marine environment.|
|The body of Hediste diversicolor may be physically damaged by mechanical interference as it has a fragile hydrostatic skeleton. Mechanical interference within the substratum, such as that caused by the dropping and dragging of an anchor or fishing gear, could physically damage ragworms within the path of the anchor and cause their displacement. Physical injury and displacement would hinder the ability of a ragworm to burrow rapidly back into the sediment to seek refuge from predation. |
Regeneration of the lost body is often observed (M. Kendall, pers. comm.) however it is likely that some individuals may die and an intolerance of intermediate has been recorded.
|Displacement from within the sediment to be left upon the sediment surface would increase the risk of Hediste diversicolor to predation but as a mobile burrowing species it is able to burrow rapidly back into the sediment and seek refuge. However, this is only possible if the animal is near its own burrow on a suitable substratum (M. Kendall, pers. comm.). The burrows of other worms are well defended through territorial behaviour.|
|Reports of the effects of synthetic chemicals on Hediste diversicolor illustrate that the intolerance of the species is highly dependent upon the molecular structure of the chemical, which determines the chemicals properties and use. For example:|
|Bryan (1984) reviewed metals in the marine environment and from the evidence available suggested that polychaetes were fairly resistant to heavy metals.|
In Hediste diversicolor the acute toxicity is dependent on the rate of uptake of the metal, since this determines the speed with which the lethal dose is built up. The rate of intake is important because this determines whether the organism's detoxification mechanisms can regulate internal concentrations. The resistance of Hediste diversicolor is thought to be dependent on a complexing system which detoxifies the metal and stores it in the epidermis and nephridia (Bryan & Hummerstone, 1971; McLusky et al. 1986).
Hediste diversicolor has been found successfully living in estuarine sediments contaminated with copper ranging from 20 µm Cu/g in low copper areas to >4000 µm Cu/g where mining pollution is encountered e.g. Restronguet Creek, Fal Estuary, Cornwall (Bryan & Hummerstone, 1971). Attempts to change the tolerance of different populations of Hediste diversicolor to different sediment concentrations of copper have shown that it is not readily achieved suggesting that increased tolerance to copper has a genetic basis (Bryan & Hummerstone, 1971; Bryan & Gibbs, 1983).
Crompton (1997) reviewed the toxic effect concentrations of metals to marine invertebrates (see Table 5.12, Crompton, 1997). Annelid species, such as Hediste diversicolor were found to be at risk if metals exceeded the following concentrations during 4-14 days of exposure: >0.1 mg Hg l-1, > 0.01 mg Cu l-1, > 1 mg Cd l-1, >1 mg Zn l-1,>0.1 mg Pb l-1, >1 mg Cr L-1, >1 mg As l-1 and >10 mg Ni l-1.
In general, for estuarine animals heavy metal toxicity increases as salinity decreases and temperature increases (McLusky et al., 1986). For example, Fernandez & Jones (1990) calculated 96 hour LC50 Zinc values for Hediste diversicolor at four salinities 5, 10, 17.5 and 30 psu at 12°C. The 96 hour LC50 at 17.5 psu and 12°C was 38 mg Zn l-1, while at 5 and 10 psu it was 7 and 19 mg Zn l-1 respectively. Toxicity decreased with increasing salinity. When salinity remained constant at 17.5 psu, but temperature varied, the following 96 hour LC 50 values for Zinc were recorded: 40 mg Zn l-1 at 6°C, 32 mg Zn l-1 at 12°C and 9.1mg Zn l-1 at 20°C. Toxicity increased with increasing temperature. Accumulation of zinc was also greater at the lowest salinities and when the temperature was highest at 20°C. In a parallel experiment, the presence of sediment was found to reduce toxicity and body accumulation of zinc in Hediste diversicolor.
Recovery of this species would be influenced by the length of time it would take for the potential habitat to return to a suitable state (e.g. factors such as the decline of bioavailable metals within the marine environment), recolonization by adult and juvenile specimens from adjacent habitats, and the establishment of a breeding population. Since juveniles remain in the infauna throughout their development selection for metal tolerance can be expected to be operative from an early stage (Bryan & Gibbs, 1983).
|The 1969 West Falmouth (America) spill of Grade 2 diesel fuel documents the effects of hydrocarbons in a sheltered habitat (Suchanek, 1993). The entire benthic fauna including Hediste diversicolor was eradicated immediately following the spill and remobilization of oil that continued for a period > 1 year after the spill, contributed to much greater impact upon the habitat than that caused by the initial spill. Effects are likely to be prolonged as hydrocarbons incorporated within the sediment by bioturbation will remain for a long time owing to slow degradation under anoxic conditions. Oil covering the surface and within the sediment will prevent oxygen transport to the infauna and promote anoxia as the infauna utilize oxygen during respiration. Although Hediste diversicolor is tolerant of hypoxia and periods of anoxia, a prolonged absence of oxygen will result in the death of it and other infauna. McLusky (1982) found that petrochemical effluents released from a point source to an estuarine intertidal mudflat, 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, and Hediste diversicolor was found amongst an impoverished fauna at 250 m from the discharge.|
|No information||No information||No information||Not relevant|
|Beasley & Fowler (1976) and Germain et al., (1984) examined the accumulation and transfers of radionuclides in Hediste diversicolor from sediments contaminated with americium and plutonium derived from nuclear weapons testing and the release of liquid effluent from a nuclear processing plant. Both concluded that the uptake of radionuclides by Hediste diversicolor was small. Beasley & Fowler (1976) found that Hediste diversicolor accumulated only 0.05% of the concentration of radionuclides found in the sediment. Both also considered that the predominant contamination pathway for Hediste diversicolor was from the interstitial water. However, there is insufficient information available on the biological effects of radionuclides to comment further upon the intolerance of this species to radionuclide contamination.|
|Nutrient enrichment favours the growth of opportunistic green macro-algae blooms which can cause declines in some species and increases in others (Raffaelli, 2000). Evidence (Beukema, 1989; Reise et al., 1989; Jensen, 1992) suggested a doubling in the abundance of Hediste diversicolor in the Dutch Wadden Sea, accompanied by a more frequent occurrence of algal blooms that were attributed to marine eutrophication. Algae may be utilized by Hediste diversicolor in its omnivorous diet, so some effects of nutrient enrichment may be beneficial to this species.|
|Hediste diversicolor is an euryhaline species, able to tolerate a range of salinities from full sea water down to 5 psu or less (Barnes, 1994). Consequently a change of one category from the MNCR salinity scale (see benchmark) for a duration of one year would not be restrictive to adults of Hediste diversicolor. Specimens already at the extreme ends of their salinity tolerance would be more intolerant of a short term change of two categories on the MNCR salinity scale but are sufficiently mobile to retreat to more hospitable conditions. Low salinities (< 8 psu) can have an adverse effect on reproduction (Ozoh & Jones, 1990; Smith 1964) (see larval sensitivity).|
|The littoral muds and muddy sands which Hediste diversicolor inhabits tend to have lower oxygen levels than other sediments. Hediste diversicolor is resistant to moderate hypoxia (Diaz & Rosenberg, 1995). The successful survival of this species under prolonged hypoxia was confirmed by the resistance experiments of Vismann (1990), which resulted in a mortality of only 15% during a 22 day exposure of Hediste diversicolor at 10% oxygen (ca. 2.8 mg O2 per litre). Hediste diversicolor is active at the sediment/water interface where hydrogen sulphide concentrations increase during periods of hypoxia. Vismann (1990), also demonstrated that the high tolerance of Hediste diversicolor to hypoxia in the presence of sulphide is enabled by elevated sulphide oxidation activity in the blood. Hediste diversicolor may also exhibit a behavioural response to hypoxia by leaving the sediment (Vismann, 1990) which is enhanced in the presence of sulphide. After 10 days of hypoxia (10% oxygen saturation) with sulphide (172-187 µmM) only 35% of Hediste diversicolor had left the sediment compared to 100% of Nereis virens. Laboratory experiments in the absence of sediments, found that Hediste diversicolor could survive hypoxia for more than 5 days and that it had a higher tolerance to hypoxia than Nereis virens, Nereis succinea and Nereis pelagica (Theede, 1973; Dries & Theede, 1974; Theede et al., 1973).|
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|No information||No information||No information||Not relevant|
|Populations of Hediste diversicolor are dominated by females, males may constitute up to 40% of the population but several reports suggest that the proportion of males is frequently lower (< 20%) (see Clay, 1967(c.)). The sexes are externally indistinguishable except when approaching maturation and during spawning (see reproduction and adult general biology). Consequently extraction e.g. by bait digging, of 50% of the specimens from within an area is likely to remove more females than males. A reduction in the female proportion of the population prior to spawning could reduce recruitment to the population. The mechanical action of the digging, even if the worms were not actually taken, may also cause some damage to the bodies. Recovery is dependent on the reproductive success and survival of the remaining population and colonization by adults from unaffected areas.|
|Many species in addition to Hediste diversicolor are taken from the intertidal environment for personal or commercial use as fishing bait. Techniques for extraction include hand digging, bait pumping and worm dredging (Fowler, 1999). Heiligenberg (1987) reported upon the effects of both hand and mechanical digging in the Dutch Wadden Sea. Hand and mechanical digging operating at a level to achieve a 50% reduction in Arenicola marina, caused a significant reduction in many of the common species, including Hediste diversicolor. A total of 1.9 g of other benthic animals were removed for every 1 g of Arenicola marina.|
Mechanical disturbance of the substrata will also displace Hediste diversicolor causing specimens to be susceptible to predation (see abrasion & displacement).
Recovery is dependent on the reproductive success and dispersion of the remaining population and colonization by adults from unaffected areas.
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|National (GB) importance||-||Global red list (IUCN) category||-|
|Origin||-||Date Arrived||Not relevant|
Hediste diversicolor may be used as bait by anglers and are often sold commercially. They are harvested using a fork to turn over the substrata and collected. Hediste diversicolor is also used as a food source in aquaculture (Scaps, 2002).
Armonies, W., Herre, E. & Sturm, M., 2001. Effects of the severe winter 1995 / 1996 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt. Helgoland Marine Research, 55, 170-175.
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Dales, R. P. & Kennedy, G.Y., 1954. On the diverse colours of Nereis diversicolor. Journal of the Marine Biological Association of the United Kingdom, 33, 699-708.
Dales, R. P., 1950. The reproduction and larval development of Nereis diversicolor O. F. Müller. Journal of the Marine Biological Association of the United Kingdom, 29, 321-360.
Dales, R.P., 1958. Survival of anaerobic periods by two intertidal polychaetes, Arenicola marina (L.) and Owenia fusiformis Delle Chiaje. Journal of the Marine Biological Association of the United Kingdom, 37, 521-529.
Davey, J.T. & George, C.L., 1986. Specific interactions in soft sediments: factors in the distribution of Nereis (Hediste) diversicolor in the Tamar Estuary. Ophelia, 26, 151-164.
Davey, J.T., 1994. The architecture of the burrow of Nereis diversicolor. Journal of Experimental Marine Biology and Ecology, 179, 155-129.
Davies, I.M, Gillibrand, P.A., McHenery, J.G. & Rae, G.H., 1998. Environmental risk of Ivermectin to sediment dwelling organisms. Aquaculture, 163, 29-46.
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Last Updated: 08/05/2008