Alitta succinea (pile worm)
- Summary of Invasiveness
- Taxonomic Tree
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Water Tolerances
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Environmental Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
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 InvasivenessTop of page
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 TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Annelida
- Class: Polychaeta
- Order: Aciculata
- Family: Nereididae
- Genus: Alitta
- Species: Alitta succinea
DescriptionTop of page
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.
DistributionTop of page
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 TableTop of page
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.Last updated: 10 Jan 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Greece||Present||Original citation: ERMS (European Register of Marine Species) (2006)|
|Italy||Present||Piave River; Sardegna; Valle di Gorino; Sacca di G|
|Russia||Present||Present based on regional distribution.|
|United Kingdom||Present||Original citation: ERMS (European Register of Marine Species) (2006)|
|Canada||Present||Present based on regional distribution.|
|United States||Present||Present based on regional distribution.|
|-Florida||Present||Pensacola Beach, Santa Rosa Island|
|-Georgia||Present||Colonels Island, Port of Brunswick|
|-Massachusetts||Present||Great Sippewissett salt marsh, Falmouth|
|-New York||Present||Introduced||Flax Pond|
|-Virginia||Present||Wachapreague, Accomack County; Fisherman Island, Northampton County; Lafayette River|
|-Washington||Present||Introduced||Puget Sound, Thurston County|
|Atlantic - Northwest||Present||Chesapeake Bay|
|Mediterranean and Black Sea||Present||Introduced||Black Sea, Sevastopol Bay|
|Pacific - Eastern Central||Present||Introduced||Lord Howe Island|
|Pacific - Northwest||Present||Introduced|
|Brazil||Present||Present based on regional distribution.|
|-Rio Grande do Sul||Present||Lagoa dos Patos, Patos Lagoon Estuary|
|Colombia||Present||Hooker Bay, San Andrés|
|Uruguay||Present||Solís Grande Stream Estuary|
History of Introduction and SpreadTop of page
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 IntroductionTop of page
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.
HabitatTop of page
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 ListTop of page
|Brackish||Inland saline areas||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Brackish||Inland saline areas||Secondary/tolerated habitat||Natural|
|Littoral||Coastal areas||Principal habitat||Harmful (pest or invasive)|
|Littoral||Coastal areas||Principal habitat||Natural|
|Littoral||Mangroves||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Littoral||Salt marshes||Principal habitat||Harmful (pest or invasive)|
|Littoral||Salt marshes||Principal habitat||Natural|
|Brackish||Estuaries||Principal habitat||Harmful (pest or invasive)|
|Brackish||Lagoons||Principal habitat||Harmful (pest or invasive)|
|Marine||Inshore marine||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Marine||Inshore marine||Secondary/tolerated habitat||Natural|
|Marine||Pelagic zone (offshore)||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Marine||Pelagic zone (offshore)||Secondary/tolerated habitat||Natural|
|Marine||Benthic zone||Principal habitat||Harmful (pest or invasive)|
|Marine||Benthic zone||Principal habitat||Natural|
Biology and EcologyTop of page
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.
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).
ClimateTop of page
|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 TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|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 DispersalTop of page
This species can disperse naturally and via water currents.
Smith and Carlton (1975) reported the introduction of A. succinea to San Francisco Bay with oysters (Crassostrea virginica and C. virginicus).
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 CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Environment (generally)||Positive and negative|
Environmental ImpactTop of page
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 FactorsTop of page
- Proved invasive outside its native range
- Highly adaptable to different environments
- Tolerant of shade
- Highly mobile locally
- 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
- Competition - monopolizing resources
- Highly likely to be transported internationally accidentally
- Difficult/costly to control
UsesTop of page
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 ListTop of page
Animal feed, fodder, forage
- Pollution indicator
Similarities to Other Species/ConditionsTop of page
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 ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
No specific information on means of control of A. succinea is reported.
ReferencesTop of page
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.
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.
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. http://www.int-res.com/abstracts/meps/v293/p155-164/
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.
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.
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.
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. http://www.marine.csiro.au/crimp/reports/PriorityPestsFinalreport.pdf
ISSG (IUCN SSC Invasive Species Specialist Group), 2013. Global Invasive Species Database (GISD). IUCN SSC Invasive Species Specialist Group. http://www.issg.org/database/welcome/
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. http://www.sfei.org/rmp/reports/Contaminant_Fate/05_No394_FateofContaminants.pdf
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.
Mistri M; Ghion F; Modugno S; Rossi R, 2002. Response of macrobenthic communities to an hydraulic intervention in an enclosed lagoon (Valle di Gorino, Northern Italy). J. Mar. Biol. Ass. UK, 82:771-778.
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.
National Introduced Marine Pest Information System (NIMPIS), 2013. Alitta succinea impacts and vectors. National Introduced Marine Pest Information System. http://www.marinepests.gov.au/nimpis
NOBANIS (North European Baltic Network on Invasive Alien Species), 2006. Neanthes succinea. North European, Baltic Network on Invasive Alien Species. http://www.nobanis.org/speciesInfo.asp?taxaID=7184
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.
Passamaneck Y; Halanych KM, 2006. Lophotrochozoan phylogeny assessed with LSU and SSU data: Evidence of lophophorate polyphyly. Mol. Phylogenet. Evol, 40(1):20-28.
Shalovenkov N, 2005. restoration of some parameters in the development of benthos after reduction of anthropogenous loading in the ecosystem of the Sevastopol Bay in the Black Sea. Mitigation and Adaption Strategies for Global Change, 10:105-113.
Ahrens M J, Hertz J, Lamoureux E M, Lopez G R, McElroy A E, Brownawell B J, 2001. The role of digestive surfactants in determining bioavailability of sediment-bound hydrophobic organic contaminants to 2 deposit-feeding polychaetes. Marine Ecology Progress Series. 145-157. DOI:10.3354/meps212145
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. 155-164. DOI:10.3354/meps293155
CABI, Undated. Compendium record. Wallingford, UK: CABI
CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Castaldelli G, Mantovani S, Welsh T, Rossi R, Mistri M, Fano E A, 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.
Elías R, Rivero M S, Vallarino E A, 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.
Hayes K, Sliwa C, Migus S, McEnnulty F, Dunstan P, 2005. National Priority Pests: Part II, Ranking of Australian marine pests. In: National Priority Pests: Part II, Ranking of Australian marine pests, Canberra, Australia: CSIRO Marine Research, Department of Environment and Heritage. 106 pp. http://www.marine.csiro.au/crimp/reports/PriorityPestsFinalreport.pdf
Mistri M, Ghion F, Modugno S, Rossi R, 2002. Response of macrobenthic communities to an hydraulic intervention in an enclosed lagoon (Valle di Gorino, Northern Italy). Journal of the Marine Biological Association of the United Kingdom. 771-778.
Pardo E V, Dauer D M, 2003. Particle size selection in individuals from epifaunal versus infaunal populations of the nereidid polychaete Neanthes succinea (Polychaeta: Nereididae). Hydrobiologia. 355-360.
Shalovenkov N, 2005. Restoration of some parameters in the development of benthos after reduction of anthropogenous loading in the ecosystem of the Sevastopol Bay in the Black Sea. Mitigation and Adaption Strategies for Global Change. 105-113.
Vary E, 2001. The Marine Invertebrates of Peggy's Cove - A Contribution to the Marine Invertebrate Diversity Initiative. In: The Marine Invertebrates of Peggy's Cove - A Contribution to the Marine Invertebrate Diversity Initiative, http://www.fundyforum.com/MIDI/events_and_docs/peggyscove.pdf
OrganizationsTop of page
Australia: Commonwealth Scientific and Industrial Research Organisation (CSIRO), CSIRO, Bag 10, Clayton South, Victoria, http://www.csiro.au/
ContributorsTop of page
22/05/08 Original text by:
Claire Beverley, CABI, Nosworthy Way, Wallingford, Oxon OX10 8DE, UK
Distribution MapsTop of page
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