Ragworm (Hediste diversicolor)
Distribution data supplied by the Ocean Biodiversity Information System (OBIS). To interrogate UK data visit the NBN Atlas.Map Help
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) |
Summary
Description
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.
Recorded distribution in Britain and Ireland
Widespread along all British coasts where suitable habitat and substratum exist.
Global distribution
Hediste diversicolor is widely distributed throughout north-west Europe on the Baltic Sea, North Sea and along Atlantic coasts to the Mediterranean.
Habitat
Inhabits muddy substrata in a more-or-less permanent U or J-shaped burrow that may be up to 20 cm in depth. Also occurs under stones on mud where the burrow is adjacent to the stone. Hediste diversicolor is widespread in brackish water environments throughout north-west Europe.
Depth range
IntertidalIdentifying features
- Head (prostonium) has four small eyes, two short antennae and two large biarticulate palps.
- Eversible pharynx with small conical chitinous teeth (paragnaths) on the proboscis.
- Four pairs of tentacular cirri, the dorsal cirri being longer than the ventral cirri.
- The neuropodial acicular ligules on the first two chaetigers have a finger-like post chaetal lobe.
- Cirri are much shorter than ligules on biramous parapodia.
- Notochaetae are long and taper to a fine point (homogomph spinigers) on biramous parapodia whilst neurochaetae may be long and tapering (homogomph and heterogomph spinigers) or have short blades with a hooked tip (heterogomph falcigers).
- In posterior chaetigers (about 40 in large animals), single stout falcigers replace heterogomph falcigers above the aciculum in neuropodia.
- Colour varies (see general biology) but mainly green or reddish.
- Prominent dorsal blood vessel.
- Inhabits mucous burrow.
Additional information
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.
Listed by
- none -
Biology review
Taxonomy
Level | Scientific name | Common name |
---|---|---|
Phylum | Annelida | Segmented worms e.g. ragworms, tubeworms, fanworms and spoon worms |
Class | Polychaeta | Bristleworms, e.g. ragworms, scaleworms, paddleworms, fanworms, tubeworms and spoon worms |
Order | Phyllodocida | |
Family | Nereididae | |
Genus | Hediste | |
Authority | (O.F. Müller, 1776) | |
Recent Synonyms | Nereis diversicolor (O.F. Müller, 1776)Nereis (Hediste) diversicolor (O.F. Müller, 1776) |
Biology
Parameter | Data | ||
---|---|---|---|
Typical abundance | High density | ||
Male size range | 60-120 mm | ||
Male size at maturity | 60-70 mm | ||
Female size range | 60-70 mm | ||
Female size at maturity | |||
Growth form | Vermiform segmented | ||
Growth rate | |||
Body flexibility | High (greater than 45 degrees) | ||
Mobility | Burrower | ||
Characteristic feeding method | Passive suspension feeder, Predator, Scavenger, Sub-surface deposit feeder, Surface deposit feeder | ||
Diet/food source | Omnivore | ||
Typically feeds on | Mud, sand & detritus. Phytoplankton & plankton. Other macrofauna. | ||
Sociability | Solitary | ||
Environmental position | Infaunal | ||
Dependency | Independent. | ||
Supports | None | ||
Is the species harmful? | No |
Biology information
Feeding. 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).
Filter-feeding. 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).
Deposit-feeding. 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 into a pellet that could 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.
Colour. 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).
Habitat preferences
Parameter | Data |
---|---|
Physiographic preferences | Enclosed coast or Embayment, Estuary, Ria or Voe |
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) |
Depth range | Intertidal |
Other preferences | No text entered |
Migration Pattern | Non-migratory or resident |
Habitat Information
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 a 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. The number 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. The entrance to the burrows of Hediste diversicolor are 1-2 mm wide in soft mud and are best seen when a fork is inserted into the sediment and the handle pulled towards the user (M. Kendall, pers. comm.). Short shallow depressions radiate from the opening: these are made by the anterior part of the worm's body as it searches for food around its hole, with the tail firmly anchored within the burrow. Burrow depth increases with body size (Esselink & Zwarts, 1989). Whilst feeding at the mud surface the worm is particularly prone to predation by wading birds and fish. The burrow is thus an important refuge in which to retreat. Esselink & Zwarts (1989) found a seasonal variation in the depth of burrows of Hediste diversicolor. The deepest burrows were seen in winter and it is likely that this deeper burrowing is an adaptation to escape low temperatures. Burrow depth was seen to level off at 15 cm because at this depth the worm can retreat from the reach of most deep-probing predators including curlews and oystercatchers. The individual burrow is a well-delimited territory but this territoriality is not extended outside the burrow (Scaps, 2002).
Life history
Adult characteristics
Parameter | Data |
---|---|
Reproductive type | Gonochoristic (dioecious) |
Reproductive frequency | Semelparous or 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 |
Larval characteristics
Parameter | Data |
---|---|
Larval/propagule type | - |
Larval/juvenile development | Oviparous |
Duration of larval stage | Not relevant |
Larval dispersal potential | 0 - 10 km |
Larval settlement period | Not relevant |
Life history information
Nereidae are monotelic, that is, they reproduce only once in their lifetime and then die (Olive & Garwood, 1981). Hediste diversicolor is gonochoristic (dioecious) and remains atokous throughout its life (Scaps, 2002). In summer and autumn, the sexes are externally indistinguishable being both reddish brown in colour. In any one population females are predominant, although to varying extent between localities (Dales, 1950; Clay, 1967 (c) and references therein). This observation led early workers to suggest parthenogenetic reproduction and hermaphroditism within Hediste diversicolor (Dales, 1950) but it is now acknowledged that these reproductive mechanisms are not found in Hediste diversicolor. Hediste diversicolor does not display epitoky or swarming behaviour associated with sexual reproduction like other nereid polychaetes, such as Nereis succinea and Nereis virens. The sex ratio in populations of Hediste diversicolor is heavily biased towards females. Olive & Garwood (1981) reported a ratio of females to males of approximately 4.6:1 in northeastern England.
Maturation & spawning. Colour differences between the sexes become more apparent upon maturation. Maturation and spawning are induced by a temperature rise in early spring to between 6°C and 11°C following a period of low winter temperatures. The male becomes bright green in colour. In contrast, the female appears darker green in colour which may be lacking on the ventral side. Reddish brown pigments may also still be visible in the female.
- Coelomic germ cells may be first recognizable in females that are at least 6 months old or about 7 cm in length. Eggs mature within the coelom surrounded by a loose mass of heterogenous corpuscles which the eggs gradually displace.
- Histolysis of the muscle layers and ingestion by phagocytes renders the female worm very brittle and enables the eggs to be released following the rupture of the body wall, which is achieved by writhing within the burrow. In addition to a rise in temperature, the lunar cycle imposes further synchrony on Hediste diversicolor so that spawning normally coincides with periods of the new or full moon.
- Mature males crawl around outside in search of a mature female and discharge sperm through the nephridia, directly outside her burrow. Direct contact between the sexes is not a necessity. Pheromones are of particular use in the final stages of reproduction for coordinating processes such as mate location and the synchronization of gamete release and spawning at the population level. The existence of pheromones has been demonstrated in a number of polychaete species (Bentley & Pacey, 1992). Dales (1950) supposed that owing to the low numbers of males in populations of Hediste diversicolor there may be some chemical signal detectable to the opposite sex. The observations of Bartels-Hardege & Zeeck (1990) support this supposition for the presence of chemical signals or sex pheromones, as males only released sperm outside burrows occupied by mature females.
- A period of increased activity follows the release of sperm, as the female and others in surrounding burrows, perform intense ventilation movements to draw sperm into their burrow. They may also carry sperm into the burrow using the proboscis in a feeding-like manner. Fertilized eggs remain inside the burrow protected by the female. Both sexes die shortly after spawning.
Environmental factors. Bartels-Hardege & Zeeck (1990) induced spawning in the laboratory, in specimens of Hediste diversicolor from tidal flats of the Jadebusen (North Sea), outside the normal spawning period of early spring. Temperatures were not lowered to simulate winter conditions but maintained at 16°C. Mature specimens appeared after four weeks and released gametes after a further four weeks according to a semilunar cycle. Reproduction was sustained for a period of four months. Such an extended spawning was witnessed on the Jadebusen following an unusually warm winter. Spawning occurred from February until May and was less synchronized. In contrast, the same population spawned within two months (February - March) following lower winter temperatures in another year. They concluded that not only a threshold temperature was important for synchronized spawning but the timing of the rise in temperature following winter was also a significant factor (Bartels-Hardege & Zeeck, 1990).
Age at maturity. Generally, Hediste diversicolor is reported to reach maturity between one and three years of age. Populations appear to show local characteristics in terms of spawning periods. Spawning may be limited to a short period in spring or extend over the summer. In the Thames Estuary, Dales (1950) reported specimens growing to maturity within one year, spawning in February, with some individuals surviving up to 18 months. Mettam et al. (1982), reported that Hediste diversicolor from the Severn Estuary matured rapidly in the spring and spawned at two years old. Olive & Garwood (1981), found that females in the Blyth Estuary, Northumberland, were in their second year before eggs began to appear, so most probably spawned in their third year. However, these authors also reported that spermatogenesis was only found to take about six months in the Blyth. Therefore, they suggested that there is a variable age at maturity and that this could have arisen either because the population were polymorphic in terms of their genetically determined age at maturity or if the age at maturity was variable and influenced by the environmental conditions (Olive & Garwood, 1981). Golding & Yuwono (1994) showed that, although full maturation of the gametes occurred, spawning was blocked by implanting cerebral ganglia from immature donors into the body cavity of adult hosts. In the Ythan Estuary, Scotland, Chambers & Milne (1975) witnessed two spawning peaks in the population of Hediste diversicolor, the first occurring between January and March, and another between June and August.
Sensitivity review
The MarLIN sensitivity assessment approach used below has been superseded by the MarESA (Marine Evidence-based Sensitivity Assessment) approach (see menu). The MarLIN approach was used for assessments from 1999-2010. The MarESA approach reflects the recent conservation imperatives and terminology and is used for sensitivity assessments from 2014 onwards.
Physical pressures
Use / to open/close text displayed
Intolerance | Recoverability | Sensitivity | Evidence / Confidence | |
Substratum loss [Show more]Substratum lossBenchmark. All of the substratum occupied by the species or biotope under consideration is removed. A single event is assumed for sensitivity assessment. Once the activity or event has stopped (or between regular events) suitable substratum remains or is deposited. Species or community recovery assumes that the substratum within the habitat preferences of the original species or community is present. Further details EvidenceHediste 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. | High | High | Moderate | Low |
Smothering [Show more]SmotheringBenchmark. All of the population of a species or an area of a biotope is smothered by sediment to a depth of 5 cm above the substratum for one month. Impermeable materials, such as concrete, oil, or tar, are likely to have a greater effect. Further details. EvidenceHediste 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). | Low | Immediate | Not sensitive | Moderate |
Increase in suspended sediment [Show more]Increase in suspended sedimentBenchmark. An arbitrary short-term, acute change in background suspended sediment concentration e.g., a change of 100 mg/l for one month. The resultant light attenuation effects are addressed under turbidity, and the effects of rapid settling out of suspended sediment are addressed under smothering. Further details EvidenceIncreased 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. | Tolerant* | Not relevant | Not sensitive* | Low |
Decrease in suspended sediment [Show more]Decrease in suspended sedimentBenchmark. An arbitrary short-term, acute change in background suspended sediment concentration e.g., a change of 100 mg/l for one month. The resultant light attenuation effects are addressed under turbidity, and the effects of rapid settling out of suspended sediment are addressed under smothering. Further details Evidence | No information | |||
Desiccation [Show more]Desiccation
EvidenceHediste 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. | Low | High | Low | Low |
Increase in emergence regime [Show more]Increase in emergence regimeBenchmark. A one hour change in the time covered or not covered by the sea for a period of one year. Further details EvidenceHediste 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. | Low | High | Low | Low |
Decrease in emergence regime [Show more]Decrease in emergence regimeBenchmark. A one hour change in the time covered or not covered by the sea for a period of one year. Further details Evidence | No information | |||
Increase in water flow rate [Show more]Increase in water flow rateA change of two categories in water flow rate (view glossary) for 1 year, for example, from moderately strong (1-3 knots) to very weak (negligible). Further details EvidenceHediste 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. | Intermediate | High | Low | Low |
Decrease in water flow rate [Show more]Decrease in water flow rateA change of two categories in water flow rate (view glossary) for 1 year, for example, from moderately strong (1-3 knots) to very weak (negligible). Further details Evidence | No information | |||
Increase in temperature [Show more]Increase in temperature
For intertidal species or communities, the range of temperatures includes the air temperature regime for that species or community. Further details EvidenceThe 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). | Intermediate | High | Low | Moderate |
Decrease in temperature [Show more]Decrease in temperature
For intertidal species or communities, the range of temperatures includes the air temperature regime for that species or community. Further details Evidence | No information | |||
Increase in turbidity [Show more]Increase in turbidity
EvidenceHediste 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. | Tolerant | Immediate | Not sensitive | Low |
Decrease in turbidity [Show more]Decrease in turbidity
Evidence | No information | |||
Increase in wave exposure [Show more]Increase in wave exposureA change of two ranks on the wave exposure scale (view glossary) e.g., from Exposed to Extremely exposed for a period of one year. Further details EvidenceHediste 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. | Intermediate | High | Low | Low |
Decrease in wave exposure [Show more]Decrease in wave exposureA change of two ranks on the wave exposure scale (view glossary) e.g., from Exposed to Extremely exposed for a period of one year. Further details Evidence | No information | |||
Noise [Show more]Noise
EvidenceHediste 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* | Low |
Visual presence [Show more]Visual presenceBenchmark. The continuous presence for one month of moving objects not naturally found in the marine environment (e.g., boats, machinery, and humans) within the visual envelope of the species or community under consideration. Further details EvidenceHediste 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. | Tolerant | Not relevant | Not sensitive | High |
Abrasion & physical disturbance [Show more]Abrasion & physical disturbanceBenchmark. Force equivalent to a standard scallop dredge landing on or being dragged across the organism. A single event is assumed for assessment. This factor includes mechanical interference, crushing, physical blows against, or rubbing and erosion of the organism or habitat of interest. Where trampling is relevant, the evidence and trampling intensity will be reported in the rationale. Further details. EvidenceThe 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. | Intermediate | High | Low | Low |
Displacement [Show more]DisplacementBenchmark. Removal of the organism from the substratum and displacement from its original position onto a suitable substratum. A single event is assumed for assessment. Further details EvidenceDisplacement 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. | Intermediate | Immediate | Very Low | Low |
Chemical pressures
Use [show more] / [show less] to open/close text displayed
Intolerance | Recoverability | Sensitivity | Evidence / Confidence | |
Synthetic compound contamination [Show more]Synthetic compound contaminationSensitivity is assessed against the available evidence for the effects of contaminants on the species (or closely related species at low confidence) or community of interest. For example:
The evidence used is stated in the rationale. Where the assessment can be based on a known activity then this is stated. The tolerance to contaminants of species of interest will be included in the rationale when available; together with relevant supporting material. Further details. EvidenceReports 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:
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 rate of decay of the synthetic chemical within the marine environment), 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. | High | Moderate | Moderate | High |
Heavy metal contamination [Show more]Heavy metal contaminationEvidenceBryan (1984) reviewed metals in the marine environment and from the evidence available suggested that polychaetes were fairly resistant to heavy metals. | Intermediate | Moderate | Moderate | High |
Hydrocarbon contamination [Show more]Hydrocarbon contaminationEvidenceThe 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. | Intermediate | High | Low | Moderate |
Radionuclide contamination [Show more]Radionuclide contaminationEvidenceBeasley & 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. | No information | No information | No information | Not relevant |
Changes in nutrient levels [Show more]Changes in nutrient levelsEvidenceNutrient 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. | Tolerant* | High | Not sensitive* | Moderate |
Increase in salinity [Show more]Increase in salinity
EvidenceHediste 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). | Low | High | Low | Low |
Decrease in salinity [Show more]Decrease in salinity
Evidence | No information | |||
Changes in oxygenation [Show more]Changes in oxygenationBenchmark. Exposure to a dissolved oxygen concentration of 2 mg/l for one week. Further details. EvidenceThe 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). | Intermediate | High | Low | High |
Biological pressures
Use [show more] / [show less] to open/close text displayed
Intolerance | Recoverability | Sensitivity | Evidence / Confidence | |
Introduction of microbial pathogens/parasites [Show more]Introduction of microbial pathogens/parasitesBenchmark. Sensitivity can only be assessed relative to a known, named disease, likely to cause partial loss of a species population or community. Further details. EvidenceInsufficient | No information | No information | No information | Not relevant |
Introduction of non-native species [Show more]Introduction of non-native speciesSensitivity assessed against the likely effect of the introduction of alien or non-native species in Britain or Ireland. Further details. EvidenceInsufficient | No information | No information | No information | Not relevant |
Extraction of this species [Show more]Extraction of this speciesBenchmark. Extraction removes 50% of the species or community from the area under consideration. Sensitivity will be assessed as 'intermediate'. The habitat remains intact or recovers rapidly. Any effects of the extraction process on the habitat itself are addressed under other factors, e.g. displacement, abrasion and physical disturbance, and substratum loss. Further details. EvidencePopulations 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. | Intermediate | High | Low | Low |
Extraction of other species [Show more]Extraction of other speciesBenchmark. A species that is a required host or prey for the species under consideration (and assuming that no alternative host exists) or a keystone species in a biotope is removed. Any effects of the extraction process on the habitat itself are addressed under other factors, e.g. displacement, abrasion and physical disturbance, and substratum loss. Further details. EvidenceMany 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. | Intermediate | High | Low | Moderate |
Additional information
Chemical factors. Determining the concentration of metals to which Hediste diversicolor is exposed is not easy. Differences in the substratum concentrations of metals indirectly reflect the differences in metal partitioning between different sediment constituents and interstitial water. In turn metal partitioning affects the bioavailabilty of metals to infaunal animals (Luoma & Bryan, 1982). Polychaetes such as Hediste diversicolor are able to absorb metals from ingested sediment as well as from solution in the surrounding water. Often it is difficult to determine which concentration, that of the sediment or interstitial water, is more significant in terms of exposure. Hediste diversicolor has a diversity of feeding modes which may also influence its exposure to chemical contaminants. Its omnivorous feeding behaviour would suggest that it takes whatever food dominates in the habitat. If the dominant food source is contaminated it will be an important route of exposure (Collier & Pinn, 1998; Bryan & Hummerstone, 1971). Wang et al. (1999) studied the assimilation efficiencies of trace metals (Ag, Cd, Co, Se & Zn) in a marine deposit-feeding polychaete, % Nereis succinea%, from ingested sediments. Their bioenergetic-based kinetic model indicated that most (>98%) of the Cd, Co, Se & Zn in polychaetes arise from sediment ingestion owing to the high ingestion rates of sediment ingestion in these animals. Wang et al. (1999) stated that environmental quality criteria must consider sediment as a potentially important source for metal uptake in benthic invertebrates.
Importance review
Policy/legislation
- no data -
Status
National (GB) importance | - | Global red list (IUCN) category | - |
Non-native
Parameter | Data |
---|---|
Native | Native |
Origin | - |
Date Arrived | Not relevant |
Importance information
Food source. Hediste diversicolor is an important prey item for wading birds because it is common, profitable and detectable as well as accessible (Zwarts & Esselink, 1989). Direct observation and analysis of the diet of wading birds indicates that Hediste diversicolor constitutes the main prey of about 15 waders and is the dominant prey for the avocet Recurvirostra avosetta, grey plover Pluvialis squatarola, curlew sandpiper Calidris ferruginea, bar-tailed godwit Limosa lapponica and curlew Numenius arquata (see Goss-Custard et al., 1977 and other references given in Zwarts & Esselink, 1989). Several flatfish prey on the macrobenthos of intertidal mudflats. Sole, Solea solea; dab, Limanda limanda, flounder, Platichthys flesus and plaice, Pleuronectes platessa, all include Hediste diversicolor in their diet.
Harvesting. Hediste diversicolor may be used as bait by anglers and is 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).
Community importance. Hediste diversicolor has been implicated as an infaunal species that plays a confounding role in the colonization and establishment of pioneering saltmarsh species. Hughes et al. (2000) reported that Hediste diversicolor reduced the success of Zostera noltei transplantation trials in southeast England. Plants protected from Hediste diversicolor had a higher survival rate, lower index of root damage and greater biomass than plants unprotected from the ragworm. In the laboratory, Hediste diversicolor was observed to grasp the leaves of Zostera noltei and pull them into the burrow. Hughes et al., (2000) considered that these results indicate that herbivory and disturbance by Hediste diversicolor was responsible, in part, for the restriction of the distribution of Zostera noltei and may be an important factor in limiting the success of transplantation experiments. Similarly, Emmerson (2000) demonstrated significant negative effects of Hediste diversicolor abundance on the survival of Spartina anglica seeds transplanted to sediment cores. Emmerson (2000) stated that there is a growing body of evidence indicating the importance of bioturbating infaunal invertebrates in regulating mudflat community structure (see Nowell et al. 1981; Levinton, 1995) and that their influence must be considered if the coastal defence functions of replanted saltmarshes are to be realized.
Bibliography
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.
Bachelet, G., 1987. Processus de recrutement er rôle des stades juvé d'invertébrés dans le fonctionnement des systèmes benthiques de substrat meuble en milieu intertidal estuarien. , Thèse d'éat, Université Bordeaux, France.
Barnes, R.S.K. & Head, S.M., 1977. Variation in paragnath number in some British populations of the estuarine polychaete Nereis diversicolor. Estuarine and Coastal Marine Science, 5, 771-781.
Barnes, R.S.K., 1994. The brackish-water fauna of northwestern Europe. Cambridge: Cambridge University Press.
Bartels-Hardege, H.D. & Zeeck, E., 1990. Reproductive behaviour of Nereis diversicolor (Annelida: Polychaeta). Marine Biology, 106, 409-412.
Bat, L., Gündoğdu, A., Akbulut, M., Çulha, M. & Satılmıs, H.H., 2001. Toxicity of zinc and lead to the polychaete Hediste diversicolor (Müller 1776). Turkish Journal of Marine Science, 7, 71-84.
Beasley, T.M. & Fowler, S.W., 1976. Plutonium and Americium: uptake from contaminated sediments by the polychaete Nereis diversicolor. Marine Biology, 38, 95-100.
Bentley, M.G. & Pacey, A.A., 1992. Physiological and environmental control of reproduction in polychaetes. Oceanography and Marine Biology: an Annual Review, 30, 443-481.
Beukema, J.J., 1990. Expected effects of changes in winter temperatures on benthic animals living in soft sediments in coastal North Sea areas. In Expected effects of climatic change on marine coastal ecosystems (ed. J.J. Beukema, W.J. Wolff & J.J.W.M. Brouns), pp. 83-92. Dordrecht: Kluwer Academic Publ.
Black, K.D., Fleming, S. Nickell, T.D. & Pereira, P.M.F. 1997. The effects of ivermectin, used to control sea lice on caged farmed salmonids, on infaunal polychaetes. ICES Journal of Marine Science, 54, 276-279.
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.]
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(4), 845-863. DOI https://doi.org/10.1017/S0025315400018014
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.
Chambers, M.R. & Milne, H., 1975. Life cycle and production of Nereis diversicolor O.F. Müller in the Ythan estuary, Scotland. Estuarine and Coastal Marine Science, 3, 133-144.
Chambers, S.J. & Garwood, P.R., 1992. Polychaetes from Scottish Waters. Part 3. Family Nereidae. Edinburgh: National Museums of Scotland.
Clay, E., 1967c. Literature survey of the common fauna of estuaries, 1. Cirratulus cirratusO.F. Müller. Imperial Chemical Industries Limited, Brixham Laboratory, PVM45/A/374.
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 (1), 131-147. DOI https://doi.org/10.1016/s0022-0981(98)00081-1
Craig, N.C.D. & Caunter, J.E., 1990. The effects of polydimethylsiloxane (PDMS) in sediment on the polychaete worm Nereis diversicolor. Chemosphere, 21, 751-759.
Crompton, T.R., 1997. Toxicants in the aqueous ecosystem. New York: John Wiley & Sons.
Daan, R. & Mulder, M., 1996. On the short-term and long-term impact of drilling activities in the Dutch sector of the North Sea ICES Journal of Marine Science, 53, 1036-1044.
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.
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.
Dries, R.R. & Theede, H., 1974. Sauerstoffmangelresistenz mariner Bodenvertebraten aus der West-lichen Ostsee. Marine Biology, 25, 327-233.
Eagle, G.A., 1983. The chemistry of sandy beach ecosystems - a review. In Sandy beaches as ecosystems (ed. A. McLachlan & T. Erasmus), pp. 203-224. The Hague, Netherlands: Junk.
Emmerson, M., 2000. Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica? Helgoland Marine Research, 54, 110-116.
Esnault, G., Retière, C. & Lambert, R., 1990. Food resource partitioning in a population of Nereis diversicolor (Annelida, Polychaeta) under experimental conditions. In Trophic relationships in the marine environment. Proceedings of the 24th European Marine Biology Symposium, Oban, United Kingdom (ed. M. Barnes & R.N. Gibson), pp. 453-467. Aberdeen: Aberdeen University Press.
Esselink, P. & Zwarts, L., 1989. Seasonal trend in burrow depth and tidal variation in feeding activity of Nereis diversicolor. Marine Ecology Progress Series, 56, 243-254.
Esselink, P., Van Belkum, J. & Esselink, K., 1989. The effect of organic pollution on the local distribution of Nereis diversicolor and Corophium volutator. Netherlands Journal of Sea Research, 23, 323-332.
Everson, C., 2000. Two species of lugworm. [On-line]. http://ourworld.compuserve.com/homepages/BMLss/lugs.htm, 2000-10-02
Fauchald, J. & Jumars, P.A., 1979. The diet of worms: a study of polychaete feeding guilds. Oceanography and Marine Biology: an Annual Review, 17, 193-284.
Fauchald, K., 1977. The polychaete worms. Definitions and keys to the orders, families and genera. USA: Natural History Museum of Los Angeles County.
Fernandez, T.V. & Jones, N.V., 1990. The influence of salinity and temperature on the toxicity of zinc to Nereis diversicolor. Tropical Ecology, 31, 40-46.
Fowler, S.L., 1999. Guidelines for managing the collection of bait and other shoreline animals within UK European marine sites. Natura 2000 report prepared by the Nature Conservation Bureau Ltd. for the UK Marine SACs Project, 132 pp., Peterborough: English Nature (UK Marine SACs Project)., http://www.english-nature.org.uk/uk-marine/reports/reports.htm
Germain, P., Miramand, P. & Masson, M., 1984. Experimental study of long-lived radionuclide transfers (americium, plutonium, technetium) between labelled sediments and annelidae (Nereis diversicolor, Arenicola marina). In International symposium on the behaviour of long-lived radionuclides in the marine environment, (ed. A.Cigna & C. Myttenaere), pp. 327-341. Luxembourg: Office for Official Publications of the European Communities.
Golding, D.W. & Yuwono, E., 1994. Latent capacities for gametogenetic cycling in the semelparous invertebrate Nereis. Proceedings of the National Academy of Sciences, 91, 11777-11781.
Gommez, J.L.C. & Miguez-Rodriguez, L.J., 1999. Effects of oil pollution on skeleton and tissues of Echinus esculentus L. 1758 (Echinodermata, Echinoidea) in a population of A Coruna Bay, Galicia, Spain. In Echinoderm Research 1998. Proceedings of the Fifth European Conference on Echinoderms, Milan, 7-12 September 1998, (ed. M.D.C. Carnevali & F. Bonasoro) pp. 439-447. Rotterdam: A.A. Balkema.
Goss-Custard, J.D., Jones, R.E. & Newberry, P.E., 1989. The ecology of the Wash. 1. Distribution and diet of wading birds (Charadrii). Journal of Applied Ecology, 14, 681-700.
Hailey, N., 1995. Likely impacts of oil and gas activities on the marine environment and integration of environmental considerations in licensing policy. English Nature Research Report, no 145., Peterborough: English Nature.
Harley, M. B., 1950. Occurrence of a filter-feeding mechanism in the polychaete Nereis diversicolor. Nature, 165, 734-735.
Hayward, P.J. & Ryland, J.S. (ed.) 1995b. Handbook of the marine fauna of North-West Europe. Oxford: Oxford University Press.
Heiligenberg, T. van den., 1987. Effects of mechanical and manual harvesting of lugworms Arenicola marina L. on the benthic fauna of tidal flats in the Dutch Wadden Sea. Biological Conservation, 39, 165-177.
Heip, C. & Herman, R., 1979. Production of Nereis diversicolor O.F. Müller (Polychaeta) in a shallow brackish water pond. Estuarine and Coastal Marine Science, 8, 297-305.
Hughes, R.G., Lloyd, D., Ball, L., Emson, D., 2000. The effects of the polychaete Nereis diversicolor on the distribution and transplantation success of Zostera noltii. Helgoland Marine Research, 54, 129-136.
Jones, L.A., Hiscock, K. & Connor, D.W., 2000. Marine habitat reviews. A summary of ecological requirements and sensitivity characteristics for the conservation and management of marine SACs. Joint Nature Conservation Committee, Peterborough. (UK Marine SACs Project report.). Available from: http://ukmpa.marinebiodiversity.org/uk_sacs/pdfs/marine-habitats-review.pdf
Kristensen, E., 1988. Factors influencing the distribution of nereid polychaetes in Danish coastal waters. Ophelia, 29, 127-140.
Levinton, J., 1995. Bioturbators as ecosystem engineers: control of the sediment fabric, inter-individual interactions and material fluxes. In Linking species and ecosystems, (ed. J.G. Jones & J.H. Lawton) pp. 29-36.
Lucas, F. & Bertru, G., 1997. Bacteriolysis in the gut of Nereis diversicolor (O.F. Müller) and effect of the diet. Journal of Experimental Marine Biology and Ecology, 215, 235-245.
Luoma, S.N. & Bryan, G.W., 1982. A statistical study of environmental factors controlling concentrations of heavy metals in the burrowing bivalve Scrobicularia plana and the polychaete Nereis diversicolor. Estuarine, Coastal and Shelf Science, 15, 95-108.
Marty, R. & Retière, C., 1999. Larval-to-juvenile mobility activities of a holobenthic species, Nereis diversicolor (O.F. Müller) (Polychaeta: Nereidae) - their involvement in recruitment. Bulletin of Marine Science, 65, 761-773.
Marty, R., 1997. Biologie de la reproduction et du développement de deux espèces d'annélides polychètes Nereis diversicolor (O.F. Müller) et Perinereis cultrifers Grübe. , Thèse de 3ème cycle, Université Rennes, France.
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.
Meador, J.P., Varanasi, U. & Krone, C.A., 1993. Differential sensitivity of marine infaunal amphipods to tributyltin. Marine Biology, 116, 231-239.
Mettam, C., Santhanam, V. & Havard, M.C.S., 1982. The oogenic cycle of Nereis diversicolor under natural conditions. Journal of the Marine Biological Association of the United Kingdom, 62, 637-645.
Moore, P.G., 1977a. Inorganic particulate suspensions in the sea and their effects on marine animals. Oceanography and Marine Biology: An Annual Review, 15, 225-363.
Nielsen, A.M., Eriksen, N.T., Iversen, J.J.L. & Riisgård, H.U., 1995. Feeding, growth and respiration in the polychaetes Nereis diversicolor (facultative filter-feeder) and Nereis virens (omnivorous) - a comparative study. Marine Ecology Progress Series, 125, 149-158.
Nowell, A.R.M., Jomars, P.A. & Eckman, J.E., 1981. Effects of biological activity on the entrainment of marine sediments. Marine Geology, 43, 133-153.
Olafsson, E.B., Peterson, C.H. & Ambrose, W.G. Jr., 1994. Does recruitment limitation structure populations and communities of macro-invertebrates in marine soft sediments: the relative significance of pre- and post-settlement processes. Oceanography and Marine Biology: an Annual Review, 32, 65-109
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.
Olivier, M., Desrosiers, G., Caron, A., Retière, C. & Caillou, A., 1995. Résponses comportementales des polychètes Nereis diversicolor (O.F. Müller) et Nereis virens (Sars) aux stimuli d'ordre alimentaire: utilisation de la matière organique particulaire (algues et halophytes). Canadian Journal of Zoology, 73, 2307-2317.
Ozoh, P.T.E. & Jones, N.N., 1990. Capacity adaptation of Hediste (Nereis) diversicolor embryogenesis to salinity, temperature and copper. Marine Environmental Research, 29 (3), 227-243. DOI https://doi.org/10.1016/0141-1136(90)90035-M
Pacey, A., 2000. Sperm motility in Arenicola marina (L.).[On-line] , 2000-10-02
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.
Philippart, C.J.M, 1994a. Interactions between Arenicola marina and Zostera noltii on a tidal flat in the Wadden Sea. Marine Ecology Progress Series, 111, 251-257.
Reise, K., 1979. Spatial configurations generated by motile benthic polychaetes. Helgoländer Wissenschaftliche Meeresuntersuchungen, 32, 55-72.
Riisgård, H.U., 1991. Suspension feeding in the polychaete Nereis diversicolor. Marine Ecology Progress Series, 70, 29-37.
Riisgård, H.U., 1994. Filter-feeding in the polychaete Nereis diversicolor: a review. Netherlands Journal of Aquatic Ecology, 28, 453-458.
Scaps, P., 2002. A review of the biology, ecology and potential use of the common ragworm Hediste diversicolor (O.F. Müller) (Annelida: Polychaeta). Hydrobiologia, 470, 203-218.
Shull, D.H., 1997. Mechanisms of infaunal polychaete dispersal and colonisation in an intertidal sandflat. Journal of Marine Research, 55, 153-179.
Smith, J.E., 1955. Salinity variation in interstitial water of sand at Kames Bay, Millport, with reference to the distribution of Nereis diversicolor Journal of the Marine Biological Association of the United Kingdom, 34, 33-46.
Smith, J.E., 1964. On the early development of Nereis diversicolor in different salinities. Journal of Morphology, 114, 437-464.
Smith, R.I., 1956. The ecology of the Tamar estuary. VII. Observations on the interstitial salinity of intertidal muds in the estuarine habitat of Nereis diversicolor. Journal of the Marine Biological Association of the United Kingdom, 35, 81-104.
Smith, R.I., 1977. Physiological and reproductive adaptations of Nereis diversicolor to life in the Baltic Sea and adjacent waters. In Essays on polychaetous annelids (ed. D.J. Reish & R. Fauchald), pp. 373-390. Los Angeles: University of Southern California.
Suchanek, T.H., 1993. Oil impacts on marine invertebrate populations and communities. American Zoologist, 33, 510-523. DOI https://doi.org/10.1093/icb/33.6.510
Theede, H., 1973. Comparative studies on the influence of oxygen deficiency and hydrogen sulphide on marine bottom invertebrates. Netherlands Journal of Sea Research, 7, 244-252.
Theede, H., Schaudinn, J. & Saffè, F., 1973. Ecophysiological studies on four Nereis species in the Kiel Bay. Oikos Supplementum, 15, 246-252,
Vismann, B., 1990. Sulphide detoxification and tolerance in Nereis (Hediste) diversicolor and Nereis (Neanthes) virens (Annelida: Polychaeta). Marine Ecology Progress Series, 59, 229-238.
Wang, Wen-Xiong, Stupakoff, I. & Fisher, N.S., 1999. Bioavailability of dissolved and sediment-bound metals to a deposit-feeding polychaete. Marine Ecology Progress Series, 178, 281-293.
Zwarts, L. & Esselink, P., 1989. Versatility of male curlews Numenius arquata preying upon Nereis diversicolor: deploying contrasting capture modes dependent on prey availability. Marine Ecology Progress Series, 56, 255-269.
Datasets
Bristol Regional Environmental Records Centre, 2017. BRERC species records recorded over 15 years ago. Occurrence dataset: https://doi.org/10.15468/h1ln5p accessed via GBIF.org on 2018-09-25.
Centre for Environmental Data and Recording, 2018. IBIS Project Data. Occurrence dataset: https://www.nmni.com/CEDaR/CEDaR-Centre-for-Environmental-Data-and-Recording.aspx accessed via NBNAtlas.org on 2018-09-25.
Cofnod – North Wales Environmental Information Service, 2018. Miscellaneous records held on the Cofnod database. Occurrence dataset: https://doi.org/10.15468/hcgqsi accessed via GBIF.org on 2018-09-25.
Environmental Records Information Centre North East, 2018. ERIC NE Combined dataset to 2017. Occurrence dataset: http://www.ericnortheast.org.ukl accessed via NBNAtlas.org on 2018-09-38
Fife Nature Records Centre, 2018. St Andrews BioBlitz 2014. Occurrence dataset: https://doi.org/10.15468/erweal accessed via GBIF.org on 2018-09-27.
Fife Nature Records Centre, 2018. St Andrews BioBlitz 2015. Occurrence dataset: https://doi.org/10.15468/xtrbvy accessed via GBIF.org on 2018-09-27.
Fife Nature Records Centre, 2018. St Andrews BioBlitz 2016. Occurrence dataset: https://doi.org/10.15468/146yiz accessed via GBIF.org on 2018-09-27.
Kent Wildlife Trust, 2018. Biological survey of the intertidal chalk reefs between Folkestone Warren and Kingsdown, Kent 2009-2011. Occurrence dataset: https://www.kentwildlifetrust.org.uk/ accessed via NBNAtlas.org on 2018-10-01.
Kent Wildlife Trust, 2018. Kent Wildlife Trust Shoresearch Intertidal Survey 2004 onwards. Occurrence dataset: https://www.kentwildlifetrust.org.uk/ accessed via NBNAtlas.org on 2018-10-01.
Lancashire Environment Record Network, 2018. LERN Records. Occurrence dataset: https://doi.org/10.15468/esxc9a accessed via GBIF.org on 2018-10-01.
Manx Biological Recording Partnership, 2017. Isle of Man wildlife records from 01/01/2000 to 13/02/2017. Occurrence dataset: https://doi.org/10.15468/mopwow accessed via GBIF.org on 2018-10-01.
Merseyside BioBank., 2018. Merseyside BioBank (unverified). Occurrence dataset: https://doi.org/10.15468/iou2ld accessed via GBIF.org on 2018-10-01.
National Trust, 2017. National Trust Species Records. Occurrence dataset: https://doi.org/10.15468/opc6g1 accessed via GBIF.org on 2018-10-01.
NBN (National Biodiversity Network) Atlas. Available from: https://www.nbnatlas.org.
OBIS (Ocean Biodiversity Information System), 2024. Global map of species distribution using gridded data. Available from: Ocean Biogeographic Information System. www.iobis.org. Accessed: 2024-10-08
South East Wales Biodiversity Records Centre, 2018. SEWBReC Worms (South East Wales). Occurrence dataset: https://doi.org/10.15468/5vh0w8 accessed via GBIF.org on 2018-10-02.
South East Wales Biodiversity Records Centre, 2018. Dr Mary Gillham Archive Project. Occurance dataset: http://www.sewbrec.org.uk/ accessed via NBNAtlas.org on 2018-10-02
Suffolk Biodiversity Information Service., 2017. Suffolk Biodiversity Information Service (SBIS) Dataset. Occurrence dataset: https://doi.org/10.15468/ab4vwo accessed via GBIF.org on 2018-10-02.
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Last Updated: 08/05/2008