Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Alitta succinea
(pile worm)



Alitta succinea (pile worm)


  • Last modified
  • 06 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Alitta succinea
  • Preferred Common Name
  • pile worm
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Annelida
  •       Class: Polychaeta
  •         Order: Aciculata
  • Summary of Invasiveness
  • A. succinea, the pile worm, is an aquatic sedentary polycheate annelid that can grow up to 19 cm long. It can be found in open seas, but appears to be able to tolerate a range of salinities and is common in estua...

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Epitoke form from de Spuikom, Oostende, Belgium. Length ca.40 mm. May 2007.
TitlePile worm
CaptionEpitoke form from de Spuikom, Oostende, Belgium. Length ca.40 mm. May 2007.
Copyright©Hans Hillewaert - CC BY-SA 3.0
Epitoke form from de Spuikom, Oostende, Belgium. Length ca.40 mm. May 2007.
Pile wormEpitoke form from de Spuikom, Oostende, Belgium. Length ca.40 mm. May 2007.©Hans Hillewaert - CC BY-SA 3.0


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Preferred Scientific Name

  • Alitta succinea Frey and Leuckart, 1847

Preferred Common Name

  • pile worm

Other Scientific Names

  • Neanthes oxypoda Marenzeller, 1879
  • Neanthes Perrieri Saint-Joseph, 1898
  • Neanthes succinea Hartman, 1938
  • Neanthes succinea Imajima, 1972
  • Nectoneanthes alatopalpis Wu et al., 1985
  • Nectoneanthes oxypoda Imajima, 1972
  • Nectoneanthes oxypoda Marenzeller, 1879
  • Nereis (Alitta) oxypoda Marenzeller, 1879
  • Nereis (Neanthes) succinea Fauvel, 1923
  • Nereis (Neanthes) succinea Hartman, 1945
  • Nereis alatopalpis Wesenberg-Lund, 1949
  • Nereis glandulosa Ehlers, 1868
  • Nereis limbata Ehlers, 1868
  • Nereis succinea Leuckart, 1847
  • Nereis succinea Wilson, 1984
  • Neries lamellosa Ehlers, 1868

International Common Names

  • English: clam worm; nereidid worm; pileworm; pile-worm; ragworm

Local Common Names

  • Germany: Meeresringelwurm
  • Netherlands: Ambergele zeeduizendpoot

Summary of Invasiveness

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A. succinea, the pile worm, is an aquatic sedentary polycheate annelid that can grow up to 19 cm long. It can be found in open seas, but appears to be able to tolerate a range of salinities and is common in estuaries and ports. It is considered native to parts of Europe’s Atlantic coast, but has been introduced widely around the world, most likely due to fouling on ships. Other means of dispersal may include oyster transport, ballast water and bait release. It impacts on native species through direct competition, but it is also known to alter the composition of the sediments in which it lives.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Annelida
  •             Class: Polychaeta
  •                 Order: Aciculata
  •                     Family: Nereididae
  •                         Genus: Alitta
  •                             Species: Alitta succinea


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A. succinea is a sedentary worm growing up to 19 cm in length with up to 160 segments, four pairs of tentacles, one pair of palps, one pair of antennae, with the parapodia differing in form from the head to the rear. The head area is darkly pigmented in contrast to the posterior region which is greenish-yellow or pale red, with white or dark spots along the whole length.

The following detailed taxonomic description is taken from Pests and Diseases Image Library (PaDIL). One pair of frontal antennae, one pair of conical palpostyles. Prostomium with entire anterior margin. Two pairs of eyes. Four pairs of tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 5-7. Maxillary ring of pharynx with conical paragnaths (papillae absent), paragnath counts: Area I: 1-7; Area II: 9-41; Area III: 13-47; Area IV: 15-37. Oral ring conical paragnaths present (papillae absent), Area V and VI present as distinct groups. Paragnath counts: Area V: 0-4 paragnaths (usually 1-3); VI: 4-19 in a roughly circular group; VII-VIII: 40-74 as a ventral band. Transverse dorsal lamellae absent. Ventrum of anterior chaetigers smooth.

Dorsal notopodial ligule present, markedly elongate and broader on posterior chaetigers. Prechaetal notopodial lobe present, approximately equal to length of dorsal notopodial ligule at least on anterior chaetigers thus notopodium of 3 similar sized ligules throughout all chaetigers. Dorsal cirrus mid-dorsally to sub-terminally attached to dorsal notopodial ligule on posterior chaetigers (not terminally attached). Dorsal cirri single. Neuropodial prechaetal lobe absent. Neuropodial postchaetal lobe present, at least on some anterior chaetigers, projecting strongly beyond end of acicular ligule, present throughout all chaetigers, digitiform. Ventral neuropodial ligule present, up to similar to length as acicular neuropodial ligule. Ventral cirri single.

Notochaetae: homogomph spinigers present (homogomph falcigers absent). Neurochaetae, dorsal fascicle: homogomph spinigers and heterogomph falcigers present (fused falcigers absent). Neurochaetae, ventral fascicle: heterogomph spinigers and falcigers present. One pair of elongate anal cirri.


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The native range of A. succineais thought to be along the Atlantic coast of Europe (Bakken and Wilson, 2005), though it is now present in many parts of the world. Populations in the North Sea, Mediterranean Sea and Black Sea coasts may be native or may have been introduced with the earliest shipping. Many other populations in eastern Asia, Africa, and Central and South America are considered cryptogenic by many authors, though are likely to have been introduced with the advent of intercontinental shipping. It is known to be introduced to Australia and the Pacific coast of North America and Hawaii. It is likely to be more widespread than indicated.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, NorthwestPresentDerrick and Kennedy, 1997Chesapeake Bay
Mediterranean and Black SeaPresentIntroducedMurina, 1997; Shalovenkov, 2005Black Sea, Sevastopol Bay
Pacific, Eastern CentralPresentIntroducedParsons, 2006Lord Howe Island
Pacific, NorthwestPresentIntroducedRay, 2005


TurkeyPresentMurina, 1997


NamibiaPresentIntroducedDay, 1967
South AfricaPresentIntroducedRay, 2005

North America

CanadaPresentPresent based on regional distribution.
-Nova ScotiaPresentVary, 2001Halifax
USAPresentPresent based on regional distribution.
-CaliforniaWidespreadIntroducedFong, 1987
-DelawarePresentPardo and Dauer, 2003Cape Henlopen
-FloridaPresentWeis and Weis, 1994Pensacola Beach, Santa Rosa Island
-GeorgiaPresentBakken and Wilson, 2005Colonels Island, Port of Brunswick
-MassachusettsPresentSardá et al., 1995Great Sippewissett salt marsh, Falmouth
-New YorkPresentIntroducedAhrens et al., 2001Flax Pond
-OregonPresentIntroducedUSGS, 2008
-VirginiaPresentPardo and Dauer, 2003Wachapreague, Accomack County; Fisherman Island, Northampton County; Lafayette River
-WashingtonPresentIntroducedUSGS, 2008Puget Sound, Thurston County

South America

ArgentinaPresentElías et al., 2003; Botto et al., 2005
BrazilPresentPresent based on regional distribution.
-Rio Grande do SulPresentBemvenuti, 1995Lagoa dos Patos, Patos Lagoon Estuary
ColombiaPresentHooker Bay, San Andrés
UruguayPresentSolís Grande Stream Estuary


BulgariaPresentMurina, 1997
DenmarkPresentIntroducedNOBANIS, North European Baltic Network on Invasive Alien Species; Muus, 1967
GermanyPresentBakken and Wilson, 2005Heligoland Island
GreecePresentERMS, European Register of Marine Species
ItalyPresentMaggiore et al., 2000; Mistri et al., 2002; Castaldelli et al., 2003; Magni et al., 2004; Munari and Mistri, 2006Piave River; Sardegna; Valle di Gorino; Sacca di G
RomaniaPresentMurina, 1997
Russian FederationPresentPresent based on regional distribution.
-Southern RussiaPresentMurina, 1997
SwedenPresentBakken and Wilson, 2005Blåbergshomen
UKPresentERMS, European Register of Marine Species
UkrainePresentMurina, 1997


AustraliaPresent1930Hayes et al., 2005
-VictoriaPresentIntroducedBakken and Wilson, 2005Hobsons Bay

History of Introduction and Spread

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Pardo and Dauer (2003) and Bakken and Wilson (2005) state that the known introduced range of A. succinea is along the coast of the Gulf of Mexico and on coasts off North, Central and South America, Europe, Africa, and the Black Sea. It also occurs in the Salton Sea (California, USA), the Caspian and Aral Seas, and in many parts of southern Australia. A. succinea was also found in the mangrove-fouling community at the Colombian Archipelago of San Andres and Old Province, Western Caribbean, and was abundant in Hooker Bay, Colombia (Londoño-Mesa et al., 2002). Maggiore et al. (2000) reported the presence of A. succinea in the Piave River estuary, North Italy in 1997. Spread in the Black Sea is reported by Murina and Michailova (1994). In addition, there are many other reports for e.g. Japan, Iran and Argentina and elsewhere in southern Africa, east and south-eastern Asia, South and Central America and even the Antartica peninsula. Few reliable records of dates of introduction exist, though it is thought to have been introduced to the Salton Sea in the 1930s (Kuhl and Oglesby, 1979) where it is now the dominant benthic macroinvertebrate, and to San Francisco Bay with Crassostrea virginica and C. virginicus between 1860 and 1910 (Smith and Carlton, 1975). See USGS (2008) for detailed collection records for the Pacific coast of the USA. It is likely that most of these introductions are accidental through ship fouling but dispersal may also be attributed to oyster transport, ballast water and bait release.  

Risk of Introduction

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Noting the widespread introduced range of A. succinea it can clearly by easily introduced. The main pathway is assumed to be fouling of ships but other potential vectors may include oyster transport, ballast water and bait release. Local spread can also be aided by ocean currents. It is thus highly likely to be further introduced, and all coastal areas not already invaded appear to be at risk from the introduction of A. succinea.



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A. succinea is found from the intertidal zone to subtidal regions, in mud and sand and under rocks or residing on artificial structures. It appears to be able to tolerate a range of salinity levels, living in U-shaped burrows in estuarine and marine habitats, and is commonly found in ports and marinas, and in and between mussels and oyster beds. However, in the Black Sea, A. succinea larvae were collected in open sea from depths of up to 1600 m and up to 125 km from the coast (Murina, 1997).

As a fouling species it is commonly found on the underside of ship hulls, particularly in the north-east Pacific (NIMPIS, 2013).

Habitat List

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Inland saline areas Secondary/tolerated habitat Harmful (pest or invasive)
Inland saline areas Secondary/tolerated habitat Natural
Estuaries Principal habitat Harmful (pest or invasive)
Estuaries Principal habitat Natural
Lagoons Principal habitat Harmful (pest or invasive)
Lagoons Principal habitat Natural
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural
Mangroves Secondary/tolerated habitat Harmful (pest or invasive)
Mangroves Secondary/tolerated habitat Natural
Salt marshes Principal habitat Harmful (pest or invasive)
Salt marshes Principal habitat Natural
Inshore marine Secondary/tolerated habitat Harmful (pest or invasive)
Inshore marine Secondary/tolerated habitat Natural
Pelagic zone (offshore) Secondary/tolerated habitat Harmful (pest or invasive)
Pelagic zone (offshore) Secondary/tolerated habitat Natural
Benthic zone Principal habitat Harmful (pest or invasive)
Benthic zone Principal habitat Natural

Biology and Ecology

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Although the genetics of A. succinea have not been widely reported in the literature, the species genome has been studied and is approximately twice the size of a similar species Alitta virens (Forde, 2013). Passamaneck (2006) conducted broader phylogenetic studies within the clade Lophotrochozoa.    

Reproductive Biology

A. succinea lives burrowed in sediment, in burrows in the mud and within dock-fouling communities of local marinas, commonly among masses of barnacles. Mature worms migrate to the water column to reproduce. Benthic adults metamorphose into the nektonic reproductive form called the heteronereid and these swim to the surface and swarm in large numbers, which facilitates fertilization and is activated by light levels (Hardege et al., 1990). The swimming (epitokol) stage lasts only a few days. Detwiler et al. (2002) reported that individuals die post spawning. Within approximately 36 hours, the eggs develop into small, setigerous two-segmented larvae (Carpelan, 1961 in Tiffany et al., 2002). The larvae are planktonic until they reach the 9 to 12 segment stage and then they start to settle in sediments to begin a benthic existence (Tiffany et al., 2002). Kuhl and Oglesby (1979) report further information on reproduction and survival.


The worm is a typical surface deposit-feeder, typically emerging at night to feed on detritus and plant material (Gillet et al., 2011). It has also been recorded with small amphipods and polychaetes in it gut contents (National Introduced Marine Pest Information System (NIMPIS), 2006). Further information on nutrition is found in Cammen (1980) and Fong (1987).

Environmental Requirements

A. succinea is a burrow-dwelling deposit feeder but its reproductive cycle requires water and it lives mostly at the water-sediment interface. It is very eurythermal and euryhaline, capable of tolerating a range of temperatures from 0.9-36ºC and varying levels of salinity (Gillet et al., 2011). This species has become accustomed to the increased salinity in the Salton Sea, California, USA, but it may not be able to persist when the salinity level exceeds the physiological limit of the species to osmoregulate (Detwiler et al., 2002). Larvae have also been found in the open sea and up to 1600 m deep (Murina, 1997). Effects of seasonally varying factors are reported by (Neuhoff, 1979), and they are known to be killed by cold (Poff et al., 1974).


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A - Tropical/Megathermal climate Tolerated Average temp. of coolest month > 18°C, > 1500mm precipitation annually
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
D - Continental/Microthermal climate Tolerated Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Dissolved oxygen (mg/l) 4 Optimum
Salinity (part per thousand) 5 67.5 Optimum
Water temperature (ºC temperature) Optimum 8-36 tolerated

Means of Movement and Dispersal

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This species can disperse naturally and via water currents.

Vector Transmission

Smith and Carlton (1975) reported the introduction of A. succinea to San Francisco Bay with oysters (Crassostrea virginica and C. virginicus).

Accidental Introduction

The main cause for long-range introductions is considered to be in ship ballast water and ship/boat hull fouling. Fouling of smaller vessels may also aid local dissemination. In addition, the use of A. succinea as a sport fishing bait, especially in the USA, could feasibly lead to further spread via unwise disposal of bait-boxes.


Pathway Causes

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CauseNotesLong DistanceLocalReferences
AquacultureOn oysters Yes Smith and Carlton, 1975
Hunting, angling, sport or racingPossibly, via sport fishing Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aquaculture stockOn oysters Yes Smith and Carlton, 1975
BaitPossibly, via sport fishing Yes
Ship ballast water and sediment Yes
Ship hull fouling Yes
WaterOcean currents Yes Yes

Impact Summary

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Environment (generally) Positive and negative

Environmental Impact

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Due to its burrowing and bottom-feeding activities, A. succinea can alter available nutrients in the sediment (Swan, 2003; Swan et al., 2007), which can affect other species living in the sediment and is thought to encourage bacterial activity (Bartoli et al., 2000). It also has an ability to aid in the transfer contaminants from the sediment to other marine organisms higher up the food chain, as it can take up and accumulate persistent trace elements and organic contaminants, for example it assimilates methyl-mercury 2-10 times more efficiently than mercury and assimilation increases when A. succinea is exposed to organic-rich sediment (Leatherbarrow et al., 2005). A. succinea was thought to have a greater impact in coastal Australia than other marine species listed (Hayes et al., 2005).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Tolerant of shade
  • Highly mobile locally
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Increases vulnerability to invasions
  • Modification of natural benthic communities
  • Modification of nutrient regime
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Fouling
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult/costly to control


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Economic Value

A. succinea is sometimes used as live bait for sport fishing.
Environmental Services

Polychaetes are one of the most useful marine organisms for detecting pollution because they reside in the water-sediment interface,and polychaetes have also been used in bioassays to monitor toxic compounds and as pollution indicators, from community or population levels to species level (Pocklington and Wells, 1992; Reish and Gerlinger, 1997). 

Uses List

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Animal feed, fodder, forage

  • Bait/attractant


  • Pollution indicator

Similarities to Other Species/Conditions

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Nereis lamellosa is very closely related to A. succinea, but it appears that few have acknowledged the differences and thus there may be some confusion between these two species in the Mediterranean area where both are present (NIMPIS, 2006). 

Prevention and Control

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No specific information on means of control of A. succinea is reported.


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Ahrens MJ; Hertz J; Lamoureux EM; Lopez GR; McElroy AE; Brownawell BJ, 2001. The role of digestive surfactants in determining bioavailability of sediment-bound hydrophobic organic contaminants to 2 deposit-feeding polychaetes. Marine Ecology Progress Series, 212:145-157.

Bakken T; Wilson RS, 2005. Phylogeny of nereidids (Polychaeta, Nereididae) with paragnaths. Zoologica Scripta, 35(5):507-547.

Bartoli M; Nizzoli D; Welsh DT; Viaroli P, 2000. Short-term influence of recolonisation by the polychaete worm Nereis succinea on oxygen and nitrogen fluxes and denitrification: a microcosm simulation. Hydrobiologia, 431:165-174.

Bemvenuti CE, 1995. The role of habitat selection and refuge in distribution and abundance of Neanthes succinea (Frey and Leuckart, 1847). Iheringia Serie Zoologia, 79:121-127.

Botto F; Valiela I; Iribarne O; Martinetto P; Alberti J, 2005. Impact of burrowing crabs on C and N sources, control, and transformations in sediments and food webs of SW Atlantic estuaries. Marine Ecology, Progress Series, 293:155-164.

Cammen LM, 1980. The significance of microbial carbon in the nutrition of the deposit feeding polychaete Nereis succinea. Marine Biology, 61(1):9-20.

Carpelan LH; Linsley RH, 1961. The pile worm, Neanthes succinea (Frey and Leuckart). The Ecology of the Salton Sea, California, in Relation to the Sportfishery. California Fish Game Fish Bulletin, 113:63-76.

Castaldelli G; Mantovani S; Welsh T; Rossi R; Mistri M; Fano EA, 2003. Impact of commercial clam harvesting on water column and sediment physicochemical characteristics and macrobenthic community structure in a lagoon (Sacca Di Goro) of the Po River Delta. Chemical and Ecology, 19(2-3):161-171.

Day JH, 1967. A Monograph on the Polychaeta of Southern Africa. Part 1. Errantia. Portsmouth, UK: Grosvenor Press.

Derrick PA; Kennedy VS, 1997. Prey selection by the hogchoker, Trinectes maculates (Pisces: Soleidae), along summer salinity gradients in Chesapeake Bay, USA. Marine Biology, 129:699-711.

Detwiler PM; Coe MF; Dexter DM, 2002. The benthic invertebrates of the Salton Sea: distribution and seasonal dynamics. Hydrobiologia, 473:139-160.

Elías R; Rivero MS; Vallarino EA, 2003. Sewage impact on the composition and distribution of polychaeta associated to intertidal mussel beds of the Mar Del Plata rocky shore, Argentina. Iheringia, Sér. Zool., Porto Alegre, 93(3):309-318.

ERMS (European Register of Marine Species), 2006. Entry for Neathes succinea. MarBEF Data System.

Fong PP, 1987. Particle-size utilization in the introduced polychaete Neanthes succinea in San Francisco Bay. Pacific Science, 41:37-43.

Forde AC, 2013. Genome size diversity and patterns within the Annelida. A thesis presented to the University of Guelph. Ontario, Canada: University of Guelph, 73 pp.

Gillet P; Surugiu F; Vasile R; Metais I; Mouloud M; Simo P, 2011. Preliminary data on population dynamics and genetics of Alitta succinea (Polychaeta: Nereididae) from the Romanian coast of the Black Sea. Italian Journal of Zoology, 78(1):229-241.

Hardege JD; Bartels-Hardege HD; Zeeck E; Grimm FT, 1990. Induction of swarming in Nereis succinea. Marine Biology, 104:291-295.

Hayes K; Sliwa C; Migus S; McEnnulty F; Dunstan P, 2005. National Priority Pests: Part II, Ranking of Australian marine pests. Canberra, Australia: CSIRO Marine Research, Department of Environment and Heritage, 106 p.

ISSG (IUCN SSC Invasive Species Specialist Group), 2013. Global Invasive Species Database (GISD). IUCN SSC Invasive Species Specialist Group.

Kuhl DL; Oglesby LC, 1979. Reproduction and survival of the pileworm Nereis succinea in higher Salton Sea salinities. Biological Bulletin, 157:153-165.

Leatherbarrow J; Ross J; David N; Yee D, 2005. Fate of contaminants in sediment of San Francisco estuary: a review of literature and data. San Francisco Estuary Institute.

Londoño-Mesa M; Polanía J; Vélez I, 2002. Polychaetes of the mangrove-fouling community at the Colombian Archipelago of San-Andres and Old Province, Western Caribbean. Wetlands Ecology and Management, 10:227-232.

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Murina VV; Michailova TV, 1994. The spread of polychaete Neanthes succinea in the benthal and pelagial of the phylophora Zerov's field in the North-West part of the Black sea. Gidrobiologiceskij Journal, 30(1):19-27.

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NOBANIS (North European Baltic Network on Invasive Alien Species), 2006. Neanthes succinea. North European, Baltic Network on Invasive Alien Species.

Pardo EV; Dauer DM, 2003. Particle size selection in individuals from epifaunal versus infaunal populations of the nereidid polychaete Neanthes succinea (Polychaeta: Nereididae). Hydrobiologia, 496:355-360.

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Swan BK; Watts JM; Reifel KM; Hurlbert SH, 2007. Role of the polychaete Neanthes succinea in phosphorus regeneration from sediments in the Salton Sea, California. Hydrobiologia, 576(1):111-125.

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Links to Websites

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National Introduced Marine Pest Information System (NIMPIS)


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Australia: Commonwealth Scientific and Industrial Research Organisation (CSIRO), CSIRO, Bag 10, Clayton South, Victoria,


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22/05/08  Original text by:

Claire Beverley, CABI, Nosworthy Way, Wallingford, Oxon OX10 8DE, UK

Distribution Maps

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