Invasion History

First Non-native North American Tidal Record:
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Marenzelleria viridis is native to the East Coast from Nova Scotia to Cape Henlopen, Delaware (Sikorski and Bick 2004) and probably Newfoundland to Chesapeake Bay; although additional sampling is needed since there are at least two additional cryptic species on the Atlantic Coast (M. neglecta and M. bastropi; Sikorski and Bick 2004; Bick 2005).

North American Invasion History:

Invasion History on the West Coast:

In 1991, worms identified as M. viridis were collected at Collinsville in the Sacramento River, California (Cohen and Carlton 1995). Their population was most consistent and abundant at this location, but they were found upstream to the Sacramento Turning Basin (Fairey et al. 2002) and downstream to Grizzly Bay and San Pablo Bay in 1992 (Cohen and Carlton 1995; Peterson and Vayssieres 2010). Marenzelleria viridis was found at the Napa River Marina, upstream from San Pablo Bay in 2004 (Cohen et al. 2005). Sikorski and Bick (2004) identify San Francisco Bay animals as M. neglecta, but note that 'all the specimens investigated were small; therefore some doubt about their identity remains’. Until molecular studies are made of the San Francisco Bay animals, we will use the name M. viridis for these populations.

Invasion History Elsewhere in the World:

Worms of the genus Marenzelleria were first reported from temperate European waters from collections made in 1979 in the Forth estuary, Scotland. Specimens collected here have been identified as M. viridis (Sikorski and Bick 2004). By the late 1980s, Marenzelleria were widespread in North Sea estuaries, and in 1985, they were collected in the Baltic Sea, at the mouth of the Oder River, Germany (Essink 1999; Bastrop and Blank 2006). By 1996, they had reached the Gulf of Bothnia and the Gulf of Finland (Leppakoski and Olenin 2000). Based on morphological and molecular surveys, M. viridis is the predominant species in North Sea estuaries (Sikorski and Bick 2004) and is common in the Baltic as far east as the Pomeranian Bight. One specimen was identified from the Gulf of Bothnia (Bastrop and Blank 2006; Blank et al. 2008). Marenzellaria neglecta was collected in the Elbe estuary, Germany (Sikorski and Bick 2004) and was abundant from the Pomeranian Bight to the Gulf of Riga in the Baltic Sea (Blank et al. 2008). From the island of Askø, Denmark north to the Gulf of Bothnia, M. arctia was the predominant species (Sikorski and Bick 2004; Bastrop and Blank 2006; Blank et al. 2008). From the 1970s to the 1990s, at least three separate contemporaneous cryptic invasions of Marenzellaria spp. into the northeastern Atlantic have occurred, probably through ballast water discharges (Sikorski and Bick 2004; Bastrop and Blank 2006).

In 2008, M. viridis was collected at one station in the Urias estuary, near Mazatlan, state of Sinaloa Mexico on the Pacific coast of Mexico (Ferrando and Mendez 2010). Since it was collected at only one station, its establishment in Pacific Mexico is uncertain.


Marenzelleria viridis is a spionid polychaete, one of several species of red-gilled mudworms. Its body consists of up to 250 setigers (setae-bearing segments). About 60-130 of these setigers bear branchiae (gills). The length of branchiae decreases over about 30-60 setigers. The body is somewhat flattened, and is dorsally convex, but ventrally somewhat concave. The body is widest anteriorly and tapers down its length. The prostomium is usually bell-shaped, rounded or bilobed anteriorly, and usually bears two pairs of eyes, deeply embedded in cuticle, in trapezoidal arrangement. The nuchal organ reaches the midsegmental ciliated crest of setiger 2. The paired tentacular palps are grooved and ciliated, spotted with black pigments, and may extend to segment 16 in fixed specimens. The parapodia begin on the 1st post-peristomial segment, and are short and biramous, with each lobe bearing a cluster of setae. The reported maximum total length ranges from 68 mm to 140 mm. The color of the body varies from green to brown, but the branchiae are always red (George 1966; Maciolek 1984; Atkins et al. 1987; Sikorski and Bick 2004; Bick 2005).


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Palpata
Order:   Canalipalpata
Suborder:   Spionida
Family:   Spionidae
Genus:   Marenzelleria
Species:   viridis


Marenzelleria jonesi (Maciolek, 1984)
Marenzelleria viridis (Maciolek, 1984)
Scolecolepides viridis (Verrill, 1873)
Scolelepides viridis (Verrill, 1873)
Scolelepis tenuis (None, None)
Scolelepis viridis (Verrill, 1873)

Potentially Misidentified Species

Marenzelleria arctia
Chamberlin 1920; Arctic Ocean, NW Pacific (Kamchatka), introduced to inner Baltic (Sikorski and Bick 2004).

Marenzelleria bastropi
Described from Currituck Sound, North Carolina (Bick 2005), full range unknown. Distinguishable primarily by molecular methods (Bick et al. 2004).

Marenzelleria neglecta
Sikorski and Bick 2004; New species, described from the Baltic Sea, considered native in North Carolina and Georgia.  They identified San Francisco Bay animals as this species, but noted doubts, since they had only young specimens (Sikorski and Bick 2004). This species has not been studied in its native Northwest Atlantic, but in the Baltic, it appears to be more confined to brackish waters. Worms that were identified as M. viridis from the North Sea coast were actually M. nelgecta (Essink 1999; Essink and Dekker 2002; Bastrop and Blank 2006).

Marenzelleria wireni
Augener 1913; Arctic Ocean coast of Alaska, Siberia (Sikorsky and Bick 2004). Worms that were identified as this species from the North Sea coast were actually M. viridis (Essink 1999; Essink and Dekker 2002; Bastrop and Blank 2006).



Marenzelleria viridis is an estuarine spionid polychaete, typically found in areas of highly variable salinity. Early accounts of its life history as 'M. cf. viridis' are somewhat variable, because they may refer to different sibling species. Descriptions of reproduction and development from Nova Scotia (George 1966), Scotland (Atkins et al. 1987), and the Netherlands ('Type II viridis', Bochert 1997) refer to M. viridis and indicate that this species spawns in spring; while an account from the Baltic (Bochert 1997, 'Type II viridis') probably refers to M. neglecta, which spawns in fall. A third species, M. arctia, of Arctic Ocean origin, is abundant in the inner Baltic (Bastrop and Blank 2006) and may have a different life history.

Marenzelleria spp. have separate sexes and mature at about 40 mm length (George 1966; Dauer et al. 1980; Bastrop and Blank 2006). Females are estimated to produce 10,000 - 46,000 eggs (George 1966; Bochert 1997). Eggs and sperm of M. viridis are shed into the water in March in Nova Scotia, Scotland, and the Netherlands (George 1966; Atkins et al. 1987; Bochert 1997). Spawning in a Baltic population (probably M. neglecta) occurs in September-December (Bochert 1997). Adults apparently die after spawning (Atkins et al. 1987). In Chesapeake Bay, adult worms with gametes were caught in plankton tows in February and March, but only on ebb tides, suggesting movement to more saline waters (Dauer et al. 1980). Eggs and early larvae of M. viridis were collected in plankton in Narragansett Bay in February and March (Paul Fofonoff, personal observations). Larvae of M. viridis from Nova Scotia settle at 10-13 setigers (George 1966), while Baltic worms settled at 15-23 setigers (Bochert 1997).

Our picture of the life history of Marenzelleria spp. is incomplete, but experimental and field data suggest that abundant populations in oligohaline and tidal fresh waters are maintained by seaward migration and spawning of adults, and the tidal transport of larvae up into estuaries (Dauer et al. 1980), a catadromous life history. George (1966) found that early larval development of M. viridis ceased at 2 PSU, and was greatly slowed at 5 PSU compared to 10 and 30 PSU. Bochert (1997) obtained similar results for a Baltic population (probably M. neglecta). However, 3-setiger larvae survive indefinitely at 2.5 PSU, while adults live well at 0.5 PSU (George 1966; Bochert 1997). These larvae could use selective tidal migration, and/or be transported up stream in saline benthic waters. The Baltic population did show decreased gametogenesis at salinities of 25-30 PSU, suggesting that M. neglecta may be a true brackish-water species, or that Baltic populations have acclimated to a low-salinity habitat (Bochert 1997).

Juveniles and adults of Marenzelleria viridis and M. neglecta inhabit mudflats and shallow muddy bottoms, usually in areas of variable or low salinity (George 1966; Atkins 1987; Peterson and Vayssieres 2010). The worms construct mucus-lined burrows, shaped like the letters J, L, or I, about 25-35 cm deep. The worm's head can protrude out of the burrow for feeding on detritus on the sediment surface, or suspended particles, caught on the palps and transported by cilia to the mouth (Dauer 1997). The animal keeps the burrow ventilated by body movements and cilia (Quintana et al. 2013; Renz and Forster 2013).


Phytoplankton; Detritus


Fishes, birds, crabs


Other polychaetes

Trophic Status:

Deposit Feeder



General HabitatTidal Fresh MarshNone
General HabitatUnstructured BottomNone
General HabitatSalt-brackish marshNone
Salinity RangeLimnetic0-0.5 PSU
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Tidal RangeSubtidalNone
Vertical HabitatEndobenthicNone
Vertical HabitatPlanktonicNone

Tolerances and Life History Parameters

Minimum Salinity (‰)0Adults frequently occur in fresh water of estuaries (George 1966
Maximum Salinity (‰)32Field, Nova Scotia (George 1966; Lab, Netherlands (Schiedek 1999)
Minimum Reproductive Temperature10George 1966, worms from Nova Scotia, development incomplete at 2 C
Minimum Reproductive Salinity10Larvae were unable to complete development at salinities below 10 ppt (George 1966)
Minimum Duration24Larval Period, 20 C- George 1966, worms from Nova Scotia
Maximum Duration40Larval Period, 10 C, 30 PSU- George 1966, worms from Nova Scotia
Minimum Length (mm)40Dauer et al. 1980
Maximum Length (mm)140Nova Scotia (George 1966)
Broad Temperature RangeNoneCold-temperate-Warm temperate
Broad Salinity RangeNoneLimnetic-Euhaline

General Impacts

Marenzelleria viridis has reached high densities in the Sacramento-San Joaquin Delta (Cohen and Carlton 1995; Peterson and Vayssieres 2010), but impacts have not been extensively studied. In European waters, M. viridis, together with M. neglecta and M. arctia, have become dominant organisms in benthic communities, partially replacing native infauna, and affecting the characteristics of sediments and their communities (Atkins et al. 1987; Hietanen et al. 2007; Hedman et al. 2011). This polychaete is a potential food source for fishes, but no economic impacts have been reported.

Ecological impacts

Competition- The invasion of M. viridis and M. neglecta in Danish and Finnish waters is associated with a sharp decline in abundance of the dominant polychaete Hediste (=Nereis) diversicolor, which may have been due to competition (Kotta et al. 2001; Delefosse et al. 2012). In the Baltic (Asko, Finland), Marenzelleria spp. displaced the native deposit-feeding amphipod Monoporeia affinis in experiments (Kotta et al. 2003). However, Marenzelleria is out-competed by the native bivalve Macoma balthica and does not successfully invade Macoma-dominated communities (Kotta et al. 2001).

Habitat Change- Changes in sediment properties and communities due to Marenzelleria spp. have been reported from European waters. The adults of M. viridis and M. neglecta create unbranched burrows down to 25-35 cm in the sediment (Atkins et al. 1987; Hietanen et al. 2007; Renz and Forster 2013). Burrow structures and sediment impacts of M. viridis and M. neglecta are similar, while M. arctia digs shallower U-shaped burrows, and has less impact on sediments (Renz and Forster 2013). Dense populations of adult worms rework the sediment, bringing buried organic materials and nutrients to the surface, possibly increasing fluxes of NH4+ and P to the water column initially, but possibly promoting phosphorus retention and nitrification in the longer term (Hietanen et al. 2007; Hedman et al. 2011; Norkko et al. 2012). Experimental studies with worms and sediments from Odense Fjord, Denmark, showed that the deeper-burrowing introduced M. viridis increased sulfur reduction and H2S in pore water, compared to the native Hediste diversicolor, favoring more sulfide-tolerant species (Kristensen et al. 2011). In the presence of Marenzelleria sp., the polychaete Hediste diversicolor (Atkins et al. 1987- Tay estuary, Scotland; Kotta et al. 2001- Baltic, Finland) and a community of oligochaetes and chironomids (Zmudzinski 1996, Vistula Lagoon, Poland) sharply declined, possibly due to competition and habitat change.

Regional Impacts

B-IINoneEcological ImpactHabitat Change
Experimental studies with worms and sediments from Odense Fjord, Denmark, showed that the deeper-burrowing introduced M. viridis increased sulfur reduction and H2S in pore water, compared to the native Nerieis diversicolor, favoring more sulfide-tolerant species (Kristensen et al. 2011).
B-XINoneEcological ImpactCompetition
Some community impacts (Zaiko et al. 2011). Marenzelleria spp. have been increasing, while the dominant amphipod, Monoporeia affinis, has decreased. This may have resulted from a decrease in primary production and food quality in the benthos. In experiments, Marenzelleria was a superior competitor when food quantity and quality were low (Eriksson Wikland and Andersson 2014).
B-VIINoneEcological ImpactCompetition
Some community impacts in Vistula Lagoon (Zaiko et al. 2011). Marenzellaria is now dominant in Vistula Lagoon infauna (Ezhova et al. 2005).
B-VIINoneEcological ImpactHabitat Change
Some habitat impacts (Zaiko et al. 2011). After the invasion of M. viridis, the benthos became more deeply distributed in the sediment, due to the deeper burrowing of M. viridis. Increased abundance of M. viridis was accompanied by a decline in corophiid amphipods, possibly due to bioturbation by the burrowing worms (Ezhova et al. 2005). Marenzelleria sp. increased oxygen penetration along its burrows, but did not effect pore water nutrient concentrations. However, at high densities, they did increase ammonium efflux. The worms did not affect total meiobenthic abundance, but did alter vertical distribution, allowing colonization deeper in the sediment (Urban-Malinga et al. 2013).
B-IVNoneEcological ImpactCompetition
A moderate level of community impacts (Zaiko et al. 2011).
B-IVNoneEcological ImpactHabitat Change
Some habitat impacts (Zaiko et al. 2011).
B-IIINoneEcological ImpactHabitat Change
Some habitat impacts (Zaiko et al. 2011), mostly due to burrowing and bioturbation.
B-XNoneEcological ImpactHabitat Change
Aeration of sediments, mitigating hypoxia (Norrko et al. 2011)
B-IINoneEcological ImpactCompetition
The invasion of M. viridis in Odense Fjord was associated with a sharp decline in abundance of the dominant polychaete Hediste diversicolor, which may have been due to competition, or to habitat change caused by the worm, or an associated increase in abundance of the lugworm Arenicola marina (Delefosse et al. 2012).
B-IIINoneEcological ImpactBioturbation
In experiments, fluxes of TCO2 and O2 increased in the presence of Marenzelleria and were higher when plant detritus was added to sediments. Marenzelleria stimulated carbon degradation and sulfate reduction by influencing pore water chemistry and dissolved organic carbon. Marenzelleria flushed out inhibitory pore water chemicals (CO2, H2S and NH4; Quintana et al. 2013; Quintana 2018).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET3 Cape Cod to Cape Hatteras 1871 Native Estab
NEA-II None 1979 Def Estab
B-XI None 1993 Def Estab
B-X None 1990 Def Estab
B-VII None 1988 Def Estab
B-V None 1985 Def Estab
B-IV None 1985 Def Estab
B-VIII None 1988 Def Estab
B-III None 2005 Def Estab
B-II None 2002 Def Estab
B-I None 2010 Def Estab
NEP-VIII None 2008 Def Unk
NA-ET2 Bay of Fundy to Cape Cod 0 Native Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 0 Native Estab
N100 Casco Bay 0 Native Estab
N130 Great Bay 0 Native Estab
N170 Massachusetts Bay 0 Native Estab
N180 Cape Cod Bay 0 Native Estab
M010 Buzzards Bay 0 Native Estab
N195 _CDA_N195 (Cape Cod) 0 Native Estab
M040 Long Island Sound 0 Native Estab
M060 Hudson River/Raritan Bay 0 Native Estab
M080 New Jersey Inland Bays 0 Native Estab
M090 Delaware Bay 0 Native Estab
M130 Chesapeake Bay 0 Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Kerfoot James R. Jr. (2022) Northward expansion leads to cold tolerance? Investigating thermal adaptation of the non?native pike killifish (Belonesox belizanus) in Florida, Environmental Biology of Fishes 105: 487–497

Atkins, S. M., Jones, A. M., Garwood, P. R. (1987) The ecology and reproductive cycle of a population of Marenzelleria viridis (Annelida: Polychaeta: Spionidae) in the Tay Estuary, Proceedings of the Royal Society of Edinburgh 92(3-4): 311-322

Bastrop, R., Rohner, M., Jurss, K. (1995) Are there two species of the polychaete genus Marenzelleria in Europe?, Marine Biology 121: 509-516

Bastrop, Ralf, Jurss, Karl, Sturmbauer, Christian (1998) Cryptic species in a marine polychaete and their independent introduction from North America to Europe, Molecular Biology and Evolution 15(2): 97-103

Bastrop, Ralf; Blank, Miriam (2006) Multiple invasions: a polychaete genus enters the Baltic Sea., Biological Invasions 8: 1195-1200

Beukema, J. J.; Dekker, R. (1995) Dynamics and growth of a recent invader into European coastal waters: the American razor clam, Ensis directus, Journal of the Marine Biological Association of the United Kingdom 75: 351-362

Beukema, J. J.; Dekker, R. (2011) Increasing species richness of the macrozoobenthic fauna on tidal flats of the Wadden Sea by local range expansion and invasion of exotic species, Helgoland Marine Research 65: 155-164

Bick, Andreas (2005) A new Spionidae (Polychaeta) from North Carolina, and a redescription of Marenzelleria wireni Augener, 1913, from Spitsbergen, with a key for all species of Marenzelleria., Helgoland Marine Research 59: 265-272

Blank, M; Laine, A. O.; Jurss, K.; Bastrop, R. (2008) Molecular identification key based on PCR/RFLP for three polychaete sibling species of the genus Marenzellaria and the species' current distribution in the Baltic Sea., Helgoland Marine Research 62: 129-141

Blank, Miriam ; Bastrop, Ralf (2009) Phylogeny of the mud worm genus Marenzelleria (Polychaeta, Spionidae) inferred from mitochondrial DNA sequences, Zoologica Scripta 38(3): 313-321

Bochert, Raft; Fritzsche, Dirk; Burckhardt, Roger (1996) Influence of salinity and temperature on growth and survival of the larvae of Marenzelleria viridis (Polychaeta, Spionidae), Journal of Plankton Research 18(7): 1239-1251

Bochert, Ralf (1997) Marenzelleria viridis (Polychaeta: Spionidae): a review of its reproduction, Aquatic Ecology 31: 163-175

Bonsdorff, Erik (2006) Zoobenthic diversity-gradients in the Baltic Sea: Continuous post-glacial succession in a stressed ecosystem., Journal of Experimental Marine Biology and Ecology 330: 383-391

Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Dauer, Daniel M. (1997) Functional morphology and feeding behavior of Marenzelleria viridis (Polychaeta: Spionidae), Bulletin of Marine Science 60(2): 512-516

Dauer, Daniel M.; Ewing, R. Michael; Tourtellotte, Gary H.; Barke, H. Russell Jr. (1980) Nocturnal swimming of Scolecolepides viridis (Polychaeta: Spionidae), Estuaries 3(2): 148-149

de Messano, Luciana Vicente Resende; Gonçalves, José Eduardo Arruda; Messano, Héctor Fabian; Campos, Sávio Henrique Calazans; Coutinho, Ricardo (2022) First report of the Asian green mussel Perna viridis (Linnaeus, 1758) in Rio de Janeiro, Brazil: a new record for the southern Atlantic Ocean , BioInvasions records 12(3): : 653–660

Delefosse, Matthieu and 6 authors (2012) Macrobenthic community response to the Marenzelleria viridis (Polychaeta) invasion of a Danish estuary, Marine Ecology Progress Series 461: 83-94

Essink, Karel (1999) Dispersal and development of Marenzelleria spp. (Polychaeta, Spionidae) populations in NW Europe and the Netherlands, Helgoländer Meeresuntersuchungen 52: 367-372

Essink, Karel; Dekkler, Rob (2002) General patterns in invasion ecology tested in the Dutch Wadden Sea: the case of a brackish-marine polychaetous worm, Biological Invasions 4: 359-368

Ezhova, Elena; Spirido, Olga (2005) Patterns of spatial and temporal distribution of the Marenzelleria cf. viridis population in the lagoon and marine environment in the southeastern Baltic Sea, Oceanological and Hydrobiological Studies 34(Suppl. 1): 209-226

Ezhova, Elena; Zmudzinski, Ludwik; Maciejewska, Krystyna (2005) Long-term trends in the macrobenthos of the Vistula Lagoon, southeastern Baltic Sea. Species composition and biomass distribution, Bulletin of the Sea Fisheries Institute (Poland) 1: 55-73

Fairey, Russell; Dunn, Roslyn; Sigala, Marco; Oliver, John (2002) Introduced aquatic species in California's coastal waters: Final Report, California Department of Fish and Game, Sacramento. Pp. <missing location>

Ferrando, Agustina; Mendez, Nuria (2010) Checklist of soft-bottom polychaetes (Annelida: Polychaeta) of the coastal lagoon Estero de Urias (Sinaloa, Mexico), Marine Biodiversity Records 3: e91

George, J. David (1966) Reproduction and early development of the spionid polychaete, Scolecolepides viridis (Verrill)., Biological Bulletin 130(1): 76-93

Gollasch, Stephan; Nehring, Stefan (2006) National checklist for aquatic alien species in Germany., Aquatic Invasions 1(2): 245-269

Granberg, Maria E.; Gunnarson, Jonas, S.; Hedman, Jenny E.; Rosenberg, Rutger; Jonsson, Per (2008) Bioturbation-driven release of organic contaminants from Baltic Sea sediments mediated by the invading polychaete Marenzelleria neglecta, Environmental Science and Technology 42: 1058-1065

Hedman, Jenny E.; Gunnarsson, Jonas S.; Samuelsson, Göran; Gilbert, Franck (2011) Particle reworking and solute transport by the sediment-living polychaetes Marenzelleria neglecta and Hediste diversicolor, Journal of Experimental Marine Biology and Ecology 407: 294-301

Hewitt, Judi E.; Norkko, Joanna; Kauppi, Laura; Villnäs, Anna; Norkko, Alf (2016) Species and functional trait turnover in response to broad-scale change and an invasive species, Ecosphere 7(3): e01289

Hietanen, Susanna; Laine, Ari O.; Lukkari, Kaarina (2007) The complex effects of the invasive polychaetes Marenzelleria spp. on benthic nutrient dynamics., Journal of Experimental Marine Biology and Ecology 352: 89-102

Holopainen, Reetta; Lehtiniemi, Maiju; Meier, H. E. Markus; Albertsson, Jan; Gorokhova, Elena; Kotta, Jonne; Viitasalo, Markku (2016) Impacts of changing climate on the non-indigenous invertebrates in the northern Baltic Sea by end of the twenty-first century, Biological Invasions Published online: <missing location>

Jaspers, Cornelia; Titelman,Josefin; Hansson, Lars Johan; Haraldsson, Matilda; Ditlefsen, Christine Røllike (2011) The invasive ctenophore Mnemiopsis leidyiiposes no direct threat to Baltic cod eggs and larvae, Limnology and Oceanography 56(2): 431-439

Jones, Nicole L.; Thompson, Janet K.; Arrigo, Kevin R.; Monismith, Stephen G. (2009) Hydrodynamic control of phytoplankton loss to the benthos in an estuarine environment, Limnology and Oceanography 54(3): 952-969

Josefson, Alf B.; Norkko, Joanna; Norkko, Alf (2012) Burial and decomposition of plant pigments in surface sediments of the Baltic Sea: role of oxygen and benthic fauna, Marine Ecological Progress Series 455: 33-49

Karlson, Agnes M. L.; Gorokhova, Elena; Elmgren, Ragnar (2015) Do deposit-feeders compete? Isotopic niche analysis of an invasion in a species-poor system, Scientific Reports 15: srep09715

Karlson, Agnes M. L.; Naslund, Johan; Ryden, Sara Blomgren; Elmgren, Ragnar (2011) Polychaete invader enhances resource utilization in a species-poor system, Oecologia 166: 1055-1065

Kotta, Jonne and 6 authors (2006) Ecological consequences of biological invasions: three invertebrate case studies in the north-eastern Baltic Sea., Helgoland Journal of Marine Research 60: 106-112

Kotta, Jonne; Orav, Helen; Sandberg-Kilpi, Eva (2001) Ecological consequence of the introduction of the polychaete Marenzelleria cf. viridis into a shallow-water biotope of the northern Baltic Sea., Journal of Sea Research 46: 273-280

Kristensen, Erik; Hansen, Tanja; Delefosse, Matthieu; Banta, Gary T. Quintana, Cintia O. (2011) Contrasting effects of the polychaetes Marenzelleria viridis and Nereis diversicolor on benthic metabolism and solute transport in sandy coastal sediment, Marine Ecology Progress Series 425: 125-139

Kube, Jan; Zettler (1996) Distribution of Marenzelleria viridis (Polychaeta: Spionidae) in the southwestern Baltic Sea in 1993/1994 - ten years after introduction., Sarsia 81: 131-142

Leidenberger, Sonja; Obst, Matthias; Kulawik, Robert; Stelzer, Kerstin; Heyer, Karin; Hardisty, Alex; Bourlat, Sarah J. (2015) Evaluating the potential of ecological niche modelling as a component in marine non-indigenous species risk assessments, Marine Pollution Bulletin 97: 470-487

Leppäkoski Erkki, Mihnea, Pia E. (1996) Enclosed seas under man-induced change: a comparsion between the Baltic and Black Seas., Ambio 25(6): 380-389

Leppakoski, Erkki; Olenin, Sergei (2000) Xenodiversity of the European brackish water seas: the North American contribution., In: Pederson, Judith(Eds.) Marine Bioinvasions. , Cambridge. Pp. 107-119

Leppakoski, Erkki; Olenin, Sergei (2000) Non-native species and rates of spread: lessons from the brackish Baltic Sea., Biological Invasions 2: 151-163

Llansó, Roberto J.; Sillett, Kristine; Scott, Lisa (2011) <missing title>, Versar, Inc., Columbia MD. Pp. <missing location>

Maciolek, Nancy J. (1984) Proceedings of the first international polychaete conference, Sydney, Australia, July 1983, Linnaean Society of New South Wales, Sydney, Australia. Pp. 48-62

Marchessaux, Guillaume; Thibault, Delphine; Claey, Cécilia (2022) An interdisciplinary assessment of the impact of invasive gelatinous zooplankton in a French Mediterranean lagoon, Biological Invasions 25(2): 499-518

Maximov, A. A. (2010) Large-scale invasion of Marenzelleria spp. Polychaeta; Spionidae) in the eastern Gulf of Finland, Baltic Sea, Russian Journal of Biological Invasions 2(1): 11-19

Neideman, Rasmus; Wenngren, Johan; Olafsson (2003) Competition between the introduced polychaeta Marenzelleria sp. andnd the native amphipod Monoporeia affinis in Baltic soft bottoms., Marine Ecology Progress Series 264: 49-55

Norrkko, Joanna and 8 authors (2012) A welcome can of worms? Hypoxia mitigation by an invasive species, Global Change Biology 18: 422-434

Olenin, Sergej, Leppakoski, Erkki (1999) Non-native animals in the Baltic Sea: alteration of benthic habitats in coastal inlets and lagoons., Hydrobiologia 393: 233-243

Orlova, Marina I.; Telesh, Irena V.; Berezina, Nadezhda A.; Antsulevich, Alexander E.; Maximov, Alexey A.; Litvinchuk, Larissa F. (2006) Effects of nonindigenous species on diversity and community functioning in the eastern Gulf of Finland (Baltic Sea)., Helgoland Journal of Marine Research 60: 98-105

Peterson, Heather A.; Vayssieres, Marc (2010) Benthic assemblage variability in the upper San Francisco estuary: A 27-year retrospective, San Francisco Estuary and Watershed Science <missing volume>: published online

Quintana, Cintia O.; Kristensen, Erik; Valdemarsen, Thomas (2013) Impact of the invasive polychaete Marenzelleria viridis on the biogeochemistry of sandy marine sediments, Biogeochemistry 115: 95-109

Quintana, Cintia Organo; Tang, Min; Kristensen, Erik (2007) Simultaneous study of particle reworking, irrigation transport and reaction rates in sediment bioturbated by the polychaetes Heteromastus and Marenzelleria, Journal of Experimental Marine Biology and Ecology 352: 392-406

Radashevsky, Vasily I. (2012b) Spionidae (Annelida) from shallow waters around the British Islands: an identification guide for the NMBAQC Scheme with an overview of spionid morphology and biology, Zootaxa 3152: 1-35

Renz, Judith R.; Forster, Stefan (2013) Are similar worms different? A comparative tracer study on bioturbation in the three sibling species Marenzelleria arctia, M. viridis,, and M. neglecta, from the Baltic Sea, Limnology and Oceanography 58(6): 2046-2058

Ristich, S. S., Crandall, M., Fortier, J. (1977) Benthic and epibenthic macroinvertebrates of the Hudson River I. Distribution, natural history, and community structure, Estuarine and Coastal Marine Science 5: 255-266

Schiedek, Doris (1999) Ecophysiological capability of Marenzelleria populations inhabiting North Sea estuaries: an overview, Helgoländer Meeresuntersuchungen 52(373-382): <missing location>

Sikorski, A. V.; Bick, A. (2004) Revision of Marenzelleria Mesnil 1898 (Spionidae, Polychaeta)., Sarsia 89: 253-275

Soors, Jan; van Haaren, Ton; Timm, Tarmo; Speybroeck, Jeroen (2013) Bratislavia dadayi (Michaelsen, 1905) (Annelida: Clitellata: Naididae): a new non-indigenous species for Europe, and other non-native annelids in the Schelde estuary, Aquatic Invasions 8: in press

Thomsen, Mads S. and 6 authors (2008) Annual changes in abundance of non-indigenous marine benthos on a very large spatial scale., Aquatic Invasions 3(2): 133-140

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database.

Urban-Malinga, Barbara; Warzocha, Jan; Zalewski, Mariusz (2013) Effects of the invasive polychaete Marenzelleria spp. on benthic processes and meiobenthos of a species-poor brackish system, Journal of Sea Research 80: 25-34

van Moorsel, Godfried; Tempelman, David; Lewis, Wilma (2010) [The polychaete worm Marenzelleria neglecta in the North Sea Canal (Polychaeta: Spionidae)], Nederlandse Faunistiche Mededelingen 34: 45-54

Wiklund, Eriksson A.-K. ; Andersson, A. (2014) Benthic competition and population dynamics of Monoporeia affinis and Marenzelleria sp. in the northern Baltic Sea, Estuarine, Coastal and Shelf Science 144: 46-53

Wittfoth, Anne K. J.; Zettler, Michael L. (2013) The application of a Biopollution Index in German Baltic estuarine and lagoon waters, Management of Biological Invasions 4: in press

Ysebaert, Tom; Meire, Patick; De Block, Marjan; De Regge, Nico; Soors, Jan (1997) A first record of Marenzellaria viridis (Verrill 1973) (Polychaeta: Spionidae) in the Schelde estuary, Belgium, Biologische Jaarboek Dodonaea 64: 176-181

Zaiko, Anastasija; Lehtiniemi, Maiju; Narscius, Aleksas; Olenin, Sergej (2011) Assessment of bioinvasion impacts on a regional scale: a comparative approach, Biological Invasions 13: 1739-1765

Zettler, Michael L.; Bick, Andreas; Bochert, Ralf (1995) Distribution and population dynamics of Marenzelleria viridis in a coastal water of the southern Baltic, Archives of Fisheries and Marine Research 42(3): 209-224

Zmudzinski, Ludwik (1996) Effect of the introduction of the American species Marenzelleria viridis (Polychaeta: Spionidae) on the benthic ecosystem of Vistula Lagoon, Marine Ecology 17(1-3): 221-226