Invasive Species Compendium

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Datasheet

Marenzelleria neglecta
(red gilled mud worm)

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Datasheet

Marenzelleria neglecta (red gilled mud worm)

Summary

  • Last modified
  • 06 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Marenzelleria neglecta
  • Preferred Common Name
  • red gilled mud worm
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Annelida
  •       Class: Polychaeta
  •         Order: Spionida
  • Summary of Invasiveness
  • The native range of M. neglecta is the Atlantic coast of the North America (Sikorski and Bick, 2004). The species is...

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Pictures

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PictureTitleCaptionCopyright
Marenzelleria neglecta; the red gilled mud worm.
TitleAdult
CaptionMarenzelleria neglecta; the red gilled mud worm.
CopyrightMerli Pärnoja
Marenzelleria neglecta; the red gilled mud worm.
AdultMarenzelleria neglecta; the red gilled mud worm.Merli Pärnoja

Identity

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Preferred Scientific Name

  • Marenzelleria neglecta Sikorski and Bick, 2004

Preferred Common Name

  • red gilled mud worm

Other Scientific Names

  • Marenzelleria cf. viridis Verrill, 1873
  • Marenzelleria Type II Bastrop et al., 1997

Local Common Names

  • Estonia: virgiina keeritsuss; virgiinia korgitsuss
  • Finland: amerikansukasmato
  • Lithuania: marenceliarija
  • Netherlands: oostzeegroenworm
  • Poland: rogoza amerykanska
  • Sweden: havsborstmask

Summary of Invasiveness

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The native range of M. neglecta is the Atlantic coast of the North America (Sikorski and Bick, 2004). The species is relatively invasive. In 1985 the non-indigenous polychaete was first found in the Baltic Sea. The potential vector of dispersal was ships’ ballast water. Within the first five years M. neglecta became established across almost the whole Baltic Sea area. Currently, the species spreads rapidly and dominates in many habitats (Bastrop and Blank, 2006). M. neglecta may also co-occur with Marenzelleria viridis, e.g. in the Elbe estuary, North Sea.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Annelida
  •             Class: Polychaeta
  •                 Order: Spionida
  •                     Family: Spionidae
  •                         Genus: Marenzelleria
  •                             Species: Marenzelleria neglecta

Notes on Taxonomy and Nomenclature

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Prior to the publication of Sikorski and Bick (2004) no distinction between Marenzelleria neglecta and Marenzelleria viridis had been made. Thus, the earlier notes on M. viridis may refer to both M. viridis and M. neglecta. These species are very similar and only large worms can be identified to the species level. However, only genetic analysis and autecological investigations confirm the taxonomical identity of the species.

Description

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M. neglecta has long worm-like greenish body with rows of short chaeta along both sides but without dorsal scales. The anterior end is with various long appendages with more than one pair of long tentacles or gills on the anterior end starting from the first setiger. There are hooked chaetae in both notopodia and neuropodia beginning far backwards of chaetal segments XX. In I or I-II some chaetae chair are considerably longer than the rest. Numerous small appendages surround anus. M. neglecta can be identified by the combination of the length of nuchal organs (up to setiger 4) and the number of branchiate setigers in relation to the total number of setigers (which is about one quarter to one third) (Sikorski and Bick, 2004).

Distribution

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M. neglecta occurs in marine and brackish waters. It inhabits vertical mucus-lined burrows (down to 40 cm in depth), feeding on sediment particles and meiobenthic and planktonic organisms (Sikorski and Bick, 2004).

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, NortheastWidespreadIntroduced1983 Invasive Essink and Kleef, 1988First recorded in Germany and the Netherlands

North America

CanadaPresentPresent based on regional distribution.
-Northwest TerritoriesWidespreadNativeSikorski and Bick, 2004Tuktoyaktuk Harbour
USAPresentPresent based on regional distribution.
-CaliforniaWidespreadNativeSikorski and Bick, 2004
-GeorgiaWidespreadNativeSikorski and Bick, 2004Ogeechee River
-MarylandWidespreadNativeSikorski and Bick, 2004
-North CarolinaWidespreadNativeSikorski and Bick, 2004Currituck Sound
-North DakotaWidespreadNativeSikorski and Bick, 2004Durham, Fox Point
-VirginiaWidespreadNativeSikorski and Bick, 2004Chester River, Chesapeake Bay

Europe

DenmarkWidespreadIntroduced Invasive NOBANIS, 2009
EstoniaWidespreadIntroduced1991 Invasive Kotta and Kotta, 1998Gulf of Riga, Vainameri Archipelago Sea, Gulf of Finland, Baltic Proper
FinlandWidespreadIntroduced1990 Invasive Norkko et al., 1993Gulf of Finland, Gulf of Bothnia
GermanyWidespreadIntroduced1983 Invasive Essink and Kleef, 1988
LatviaWidespreadIntroduced1988 Invasive Lagzdins and Pallo, 1994Baltic Proper, Gulf of Riga
LithuaniaWidespreadIntroduced1988 Invasive Olenin and Chubarova, 1992; Zmudzinski et al., 1993Curonian Lagoon, Baltic Proper
NetherlandsWidespreadIntroduced1983 Invasive Essink and Kleef, 1988
NorwayLocalisedIntroduced1990 Invasive NOBANIS, 2009
PolandWidespreadIntroduced1988 Invasive Gruszka, 1991Baltic Proper, Szczecin Lagoon, Vistula Lagoon
Russian FederationPresentPresent based on regional distribution.
-Central RussiaWidespreadIntroduced1996 Invasive Maximov and Panov, 2002Luga Bay, Neva Bay
SwedenWidespreadIntroduced1990 Invasive Jansson, 1994Baltic Proper

History of Introduction and Spread

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The first record of M. neglecta on the European coast is dated 1983. The species was caught in the Ems estuary between Germany and The Netherlands (Essink and Kleef, 1988). M. neglecta quickly invaded nearly all European intertidal and estuarine areas. In 1985 the species was recorded in the German Darb-Zingst bodden chain, Baltic Sea (Bick and Burkhardt, 1989). In a few years it was discovered in Poland (Gruszka, 1991), Lithuania (Zmudzinski et al., 1993) and the whole Gulf of Riga (both Latvian and Estonian coasts) (Lagzdins and Pallo, 1994, database of the Estonian Marine Institute). In 1990 it reached the Swedish coast (Jansson, 1994) and was observed in Tvärminne, Finland (Norkko et al., 1993).

From 1990 to 1993 M. neglecta expanded to the eastern Gulf of Finland and the southern Gulf of Bothnia with Olkiluoto (Finland) being the northernmost observation point (Stigzelius et al., 1997). In 1995 the species was found in the Väinameri Archipelago Sea (between the Estonian mainland and western islands).
 
In 1996, M.neglecta was first recorded in the easternmost Gulf of Finland near Luga Bay. In 1997 juveniles of M. neglecta were also found. In 2000 the polychaete was found along the whole southern coast of the gulf up to the freshwater Neva Bay. Recent surveys have shown that this species has already become the most common component of benthic invertebrates in the eastern Gulf of Finland (Maximov and Panov, 2002).
 
Genetic differences between populations of Marenzelleria in the North Sea and the Baltic Sea as well as differences in reproduction timing indicate that these populations represent two different species with two independent invasions into the two seas (Bastrop et al., 1995). The North Sea populations were described as Marenzelleria cf. wireni (type I), and the Baltic Sea populations as former Marenzelleria cf. viridis (type II), recently described as M. neglecta (Sikorski and Bick, 2004).

Risk of Introduction

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Earlier studies indicate that the species is very invasive and able to spread large distances and form permanent populations. During the initial stages of introduction the species is often present in high abundance. After expansion during the first decade of invasion, the density of M. neglecta declines in most habitats except for river estuaries and deeper seas. Nevertheless, the larvae of this polychaete are today the most prevalent taxa in meroplankton and have, in general, doubled the total meroplankton abundance (Kotta et al., 2008).

Habitat

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M. neglecta and M. viridis share habitats in the Atlantic, but diverge in their salinity preferences: M. viridis does not occur at salinities below 16‰, whereas M. neglecta prefers salinities between 0.5 and 10‰ (Sikorski and Bick, 2004). M. neglecta is an estuarine species which inhabits sandy and muddy sediments (Zettler, 1997). In shallower areas < 10 m) it prefers sand or gravel bottoms with higher abundance in more densely vegetated areas. Deeper down (> 10 m) M. neglecta is confined to silty clay substrate. It can cope with low oxygen levels and withstands salinity down to 0.5‰ (Fritzsche and von Oertzen, 1995). Successful reproduction however cannot take place at salinities below 5‰ (Norkko et al., 1993; Bochert et al., 1994).

M. neglecta lives in mucus-lined burrows with a maximum diameter of 2 mm and may penetrate down to 35 cm into the sediments. This spionid polychaete is classified as a selective surface deposit-feeder and a suspension-feeder (Dauer et al., 1981; Zettler et al., 1994). Faecal-pellet strings are deposited near the openings of the burrows.

Habitat List

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CategoryHabitatPresenceStatus
Brackish
Estuaries Principal habitat Natural
Estuaries Principal habitat Productive/non-natural
Lagoons Principal habitat Natural
Lagoons Principal habitat Productive/non-natural
Littoral
Coastal areas Principal habitat Harmful (pest or invasive)
Coastal areas Principal habitat Natural
Mud flats Principal habitat Natural
Mud flats Principal habitat Productive/non-natural
Marine
Inshore marine Principal habitat Natural
Inshore marine Principal habitat Productive/non-natural
Pelagic zone (offshore) Principal habitat Natural
Pelagic zone (offshore) Principal habitat Productive/non-natural

Biology and Ecology

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M. neglecta has a life span of approximately three years. M. neglecta is benthic as an adult and has pelagic larvae. Many individuals reach sexual maturity after one year. Benthic adult worms inhabit vertical mucus-lined burrows and feed on sediment particles, planctonic, meiobenthic organisms and resuspended organic material (Daunys et al., 1999).

M. neglecta is known to enhance the content of sediment chlorophyll a and reduce growth and survival of the polychaete Hediste diversicolor and growth of the amphipod Monoporeia affinis. The survival of M. neglecta was significantly reduced by the presence of Macoma balthica. Field data agrees with the experimental finding that M. affinis was not found even at moderate densities of M. neglecta. On the other hand, the negative effect of M. balthica on M. neglecta was observed in pristine conditions and not in eutrophicated areas (Kotta et al., 2004; 2006).
 
Genetics
 
M. neglecta is particularly difficult to distinguish from M. viridis. Both species may co-occur and only genetic analysis may confirm exact identity of the species (Bick and Zettler, 1997; Sikorski and Bick, 2004).
 
Reproductive Biology
 
Gametogenesis of M. neglecta starts in spring. Fecundity of animals depends on salinity, temperature, age and body size. Development of gametes starts in mid-May. The individuals reach maturity in September after about 20 weeks. Animals spawn in autumn and sometimes larval abundance reaches up to 21´106 individuals/m-3 near the coast.
 
The pelagic larvae can be found mainly from September to November, but may also occur up to March (Bochert, 1997; Sikorski and Bick, 2004). Reproductive features of the Baltic M. neglecta differ from M. wireni (and M. viridis) populations from the North Sea and North America. The two types reproduce at different times; M. wireni reproduces in spring while M. neglecta reproduces in autumn (Bastrop et al., 1995; Bochert, 1997). In the Gulf of Riga, however, the maximum peak of larvae is recorded in spring (Kotta et al., 2006; database of the Estonian Marine Institute). Both types show differences in larval development, gametal development and sex ratio of mature worms (Bastrop et al., 1995; Bochert, 1997).
 
Larval development largely depends on water temperature and lasts about 4 to 12 weeks. The larvae have initially one and later two pairs of black eyes and, in the 2nd segment, a ciliated pit. Neuropodial setae develop once the 7-setiger stage is reached. Palps appear at the 10-setiger stage. Metamorphosis into the juvenile benthic stage takes place at the earliest when the 15-setiger stage has been reached. Successful larval development from egg to juvenile is not possible below salinities of 5 psu, but colonization of oligohaline regions can be accomplished by larvae with more than 4 setigers or by swimming juveniles.
           
Physiology and Phenology
 
M. neglecta prefers salinities between 0.5 and 10‰ (Sikorski and Bick, 2004). It can cope with low oxygen levels and withstands salinity down to 0.5‰ (Fritzsche and von Oertzen, 1995). Successful reproduction however cannot take place at salinities below 5‰ (Norkko et al., 1993; Bochert et al., 1994).
 
Nutrition
 
M. neglecta tolerates and even prefers eutrophicated conditions. In such habitats the species can be found at high numbers and it may outcompete native species.
 
Associations
 
M. neglecta inhabits soft sediments in association with large number of benthic invertebrates such as clams, bivalves, amphipods and other polychaete worms. However, the species is not directly dependent on these associate species.
 
Environmental Requirements
 
See notes on Habitat and Physiology and Phenology.

Climate

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ClimateStatusDescriptionRemark
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Ds - Continental climate with dry summer Preferred Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) 0.5 30 Optimum 0-92 tolerated (Olenin and Chubarov, 1994; Norkko et al., 1995; Database of the Estonian Marine Institute)
Salinity (part per thousand) 5 10 Optimum 0.5-10 tolerated although cannot reproduce at salinities <5

Notes on Natural Enemies

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Several benthophagous fish species feed on M. neglecta (Olenin, 2006). The survival of M. neglecta is significantly reduced by the presence of Macoma balthica (Kotta et al., 2004). Field data agrees with experimental findings that although the negative effect of M. balthica on M. neglecta was observed in pristine conditions, it was not observed in eutrophicated areas (Kotta et al., 2006).

Competitive interactions between M. neglecta and native fauna may explain why the polychaete densities are low when the densities of M. balthica are high. Competitive superiority of M. balthica over M. neglecta is likely to be due to this species’ efficient feeding modes and better tolerance of food shortages (e.g. Brafield and Newell, 1961; Ólafsson, 1986; Kotta et al., 2004). Because M. balthica is one of the most common species in the soft bottom sediments (Segerstråle, 1957; Hällfors et al., 1981), competitive interactions between M. neglecta and M. balthica appear to be a key factor limiting the further expansion of M. neglecta in the study area.

Means of Movement and Dispersal

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Distribution of the genus Marenzelleria is restricted to the northern hemisphere. The native range of M. neglecta is the Atlantic coast of the North America (Sikorski and Bick, 2004). Within its native range and invaded habitats the species disperses by means of larvae. Ballast water is the most likely vector of its intercontinental dispersal.

Natural Dispersal (Non-Biotic)

M. neglecta is benthic as an adult and has pelagic larvae. The distribution mainly takes place via larval development when the individuals can be transported by currents over hundreds of km.

Accidental Introduction

The species has been introduced to the Baltic and North Seas by means of shipping (ballast water) (Bastrop et al., 1998). The species was most likely transported as larvae and/or adults in ballast water. This assumption is supported by collection of specimens on a plankton tow in a North American estuary (Maciolek, 1984). Shipping may influence the spread of M. neglecta although its spread is mostly associated with dispersal and development of planktonic larvae which can also be transported in ballast waters of ships.

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Ship ballast water and sedimentplanktonic larvae, benthic adults Yes

Impact Summary

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CategoryImpact
Economic/livelihood Negative
Environment (generally) Negative

Impact

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The species has a moderate impact on the recipient communities both in shallow and deep water habitats; however, severe economic and social impacts are not known.

Environmental Impact

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M. neglecta is known to enhance the content of sediment chlorophyll a and reduce growth and survival of the polychaete Hediste diversicolor and growth of the amphipod Monoporeia affinis. The survival of M. neglecta was significantly reduced by the presence of Macoma balthica. Field data agrees with the experimental findings that M. affinis was not found even at moderate densities of M. neglecta, and M. affinis has not recovered in areas with a high density of M. neglecta. However, the negative effects of M. balthica on M. neglecta was only observed in pristine conditions and not in eutrophicated areas (Kotta et al., 2004; 2006).

Higher sediment chlorophyll a content in the treatments with M. neglecta indicates higher biodeposition and/or bioturbating activity of the polychaete as compared to the native fauna. According to Pelegri and Blackburn (1995), polychatetes significantly accelerate nitrogen remineralisation and transformation processes within the sediment. As M. neglecta burrows much deeper and more actively than the native polychaetes the amount of reworked sediments, i.e. the availability of nutrients to microalgal growth, is higher in the presence of M. neglecta.

Other known effects are the increased benthic production and higher burrowing activity that improves oxygen circulation in the sediment. The introduction of the species has resulted in changes of the diet of benthic fish (Database of the Estonian Marine Institute).

Impact on Habitats

M. neglecta competes with native benthic macrofauna for food and space. Being numerically dominant it can change the structure of a native benthic community. The burrowing activity of this worm has a high impact on fluid-exchange rates between bottom water and sediments, especially in muddy sediments. The burrow walls make good substrates for aerobic degradation of organic matter (HELCOM, 1996; Olenin, 2006).

Impact on Biodiversity

M. neglecta is a deep burrowing deposit feeder and thus represents a new function in some recipient areas (e.g. the northern Baltic Sea). Experimental studies have shown that M. neglecta negatively influence the native polychaete species Hediste diversicolor and the amphipod Monoporeia affinis. Recent observational studies also demonstrate an inverse abundance and biomass relationship between M. neglecta and H. diversicolor (Kotta et al., 2004). This indicates that the species may strongly affect the composition of native benthic communities and influence the balance of organisms in a particular ecosystem (Kotta et al., 2001; Kotta and Olafsson, 2003; Kotta et al., 2004). However, the negative effects are likely to decrease with the increasing density of the adult specimens of the native bivalve Macoma balthica. Competitive superiority of M. balthica over M. neglecta is likely due to the more efficient feeding regime of the bivalve (Kotta et al., 2004; Kotta, 2005).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Pioneering in disturbed areas
  • Highly mobile locally
  • Fast growing
  • Has high reproductive potential
Impact outcomes
  • Altered trophic level
  • Conflict
  • Ecosystem change/ habitat alteration
  • Modification of natural benthic communities
  • Modification of nutrient regime
  • Modification of successional patterns
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - smothering
  • Filtration
  • Predation
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control

Uses List

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Animal feed, fodder, forage

  • Fishmeal

Detection and Inspection

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Standard benthic invertebrate sampling methods (e.g. using various benthic grabs or corers) may be used. Specific methods not known.

Similarities to Other Species/Conditions

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M. neglecta is particularly difficult to distinguish from M. viridis. Both species may co-occur (e.g. in the Elbe estuary, North Sea) and only large worms can be reliably identified. Only genetic analysis and autecological investigations may provide solutions to this problem (Bick and Zettler, 1997; Sikorski and Bick, 2004).

Prevention and Control

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Prevention

Specific methods not known. Ballast water treatment provides a good solution to reduce the invasion of the species into new recipient areas.
 
Control
 
Efficient methods of control are not known.
 
Monitoring and Surveillance
 
Standard benthic invertebrate sampling methods (e.g. using various benthic grabs or corers) may be used. Specific methods not known.
 
Ecosystem Restoration
 
Not known. However, the impacts are not so severe that requires ecosystem restoration.
 

Gaps in Knowledge/Research Needs

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There is a need for better taxonomic understanding of the species.

 

References

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Aquatic alien species in German inland and coastal waters, 2009. www.aquatic-aliens.de. http://www.aquatic-aliens.de/species-directory.htm

Bastrop R; Blank B, 2006. Multiple invasions - a polychaete genus enters the Baltic Sea. Biological Invasions, 8(5):1195-1200.

Bastrop R; Jürss K; Sturmbauer C, 1998. Cryptic species in a marine polychaeta and their independent introduction from North America to Europe. Molecular Biology and Evolution, 15:97-103.

Bastrop R; Röhner M; Jürss K, 1995. Are there two species of the polychaete genus Marenzelleria in Europe? Marine Biology, 121:509-516.

Bastrop R; Röhner M; Sturmbauer C; Jürss K, 1997. Where did Marenzelleria spp. (Polychaeta: Spionidae) in Europe come from? Aquatic Ecology, 31(2):119-136.

Bick A; Burckhardt R, 1989. First record of Marenzelleria viridis (Polychaeta, Spionidae) in the Baltic Sea, with a key to the Spionidae of the Baltic Sea. Mitteilungen aus dem Zoologischen Museum in Berlin, 65:237-247.

Bick A; Zettler ML, 1997. On the identity and distribution of two species of Marenzelleria (Polychaeta: Spionidae) in Europe and North America. Aquatic Ecology, 3(2):137-148.

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

Bochert R; Bick A; Zettler M; Arndt E-A, 1994. Marenzelleria viridis (Verrill, 1873) (Polychaeta, Spionidae), an invader in the benthic community in Baltic coastal inlets - investigations of reproduction. In: Proceeding of 13th Symposium of the Baltic Marine Biologists, Riga, Latvia.

Brafield AE; Newell GE, 1961. The behavior of Macoma baltica (L.). Journal of the Marine Biological Association of the United Kingdom, 44:81-87.

Dauer DM; Maybury CM; Ewing RM, 1981. Feeding behavior and general ecology of several spionid polychaetes from the Chesapeake Bay. Journal of Experimental Marine Biology and Ecology, 54:21-38.

Daunys D; Zettler ML; Gollasch S, 1999. Marenzelleria cf. viridis (Verrill, 1873) Annelida, Polychaeta, Spionidae. Exotics across the ocean. In: Case histories on introduced species [ed. by Gollasch S, Minchin D, Rosental H, Voigt M] University of Kiel, Germany: Verlag, 61-67.

Essink K; Kleef HL, 1988. Marenzelleria viridis (Verrill, 1873) (Polychaeta: Spionidae): a new record from the Ems estuary (The Netherlands/Federal Republic of Germany). Zoologische Bijdragen, 38:1-13.

Fritzsche D; Oertzen JAvon, 1995. Metabolic responses to changing environmental conditions in the brackish water - polychaetes Marenzelleria viridis and Hediste diversicolor. Marine Biology, 121:693-699.

Gruszka P, 1991. Marenzelleria viridis (Verrill, 1873) (Polychaeta, Spionidae) - a new component of shallow water benthic community in the Southern Baltic. In: Proc. 11th Baltic Mar. Biol. Symposium, Szczecin, Poland, 1989, 65.

HELCOM, 1996. 3rd Periodic Assessment of the State of the Marine Environment of the Baltic Sea.

Hällfors G; Niemi A; Ackefors H; Lassig J; Leppäkoski E, 1981. Biological oceanography. In: The Baltic Sea [ed. by Voipio A] Amsterdam, Netherlands: Elsevier Oceanography, 219-274.

Jansson K, 1994. Alien species in the marine environment. Introductions to the Baltic Sea and the Swedish west coast. Report 4357. Solna, Sweden: Swedish Environmental Protection Agency, 59-68.

Kotta J, 2005. Estonian Marine Institute, University of Tartu -Fact sheet on Marenzelleria cf. Viridis. http://www.sea.ee/Sektorid/merebioloogia/MASE/Benthic_invertebrates.htm

Kotta J; Kotta I, 1998. Distribution and invasion ecology of Marenzelleria viridis (Verrill) in the Estonian coastal waters. Proceedings of the Estonian Academy of Sciences. Biology. Ecology, 47:210-217.

Kotta J; Kotta I; Simm M; Lankov A; Lauringson V; Põllumäe A; Ojaveer H, 2006. Ecological consequences of biological invasions: three invertebrate case studies in the north-eastern Baltic Sea. Helgoland Marine Research, 60(2):106-112. http://www.springerlink.com/(vbc5i445bmwdkp45y4ynpj45)/app/home/contribution.asp?referrer=parent&backto=issue,6,13;journal,2,169;linkingpublicationresults,1:103796,1

Kotta J; Lauringson V; Martin G; Simm M; Kotta I; Herkül K; Ojaveer H, 2008. Gulf of Riga and Pärnu Bay. In: Ecology of Baltic Coastal waters [ed. by Schiewer U] Heidleberg, Germany: Springer, 217-243.

Kotta J; Ólafsson E, 2003. Competition for food between the introduced exotic polychaete Marenzelleria viridis and the resident native amphipod Monoporeia affinis in the Baltic Sea. Journal of Sea Research, 50(1):27-35.

Kotta J; Orav H; Sandberg-Kilpi E, 2001. Ecological consequence of the introduction of the polychaete Marenzelleria viridis into a shallow water biotope of the northern Baltic Sea. Journal of Sea Research, 46:273-280.

Kotta J; Orav-Kotta H; Sandberg-Kilpi E, 2004. Changes in the feeding behaviour of benthic invertebrates: effect of the introduced polychaete Marenzelleria viridis on the Baltic clam Macoma balthica. Proceedings of the Estonian Academy of Sciences. Biology. Ecology, 53:269-275.

Lagzdins G; Pallo P, 1994. Marenzelleria viridis (Verrill) (Polychaeta, Spionidae) - a new species for the Gulf of Riga. Proceedings of the Estonian Academy of Sciences Biology, 43:184-188.

Maciolek NJ, 1984. New records and species of Marenzelleria Mesnil and Scolecolepides Ehlers (Polychaeta: Spionidae) from north-eastern North America. In: Proceedings of the first international Polychaete Conference, Australia, Sydney [ed. by Hutchings PA], 48-62.

Maximov AA; Panov VE, 2002. Expansion of Marenzelleria viridis (Polychaeta: Spionidae) in the eastern Gulf of Finland. In: The Gulf of Finland Symposium, Estonia, Tallinn, 28-30 October 2002.

NOBANIS, 2009. Factsheet on Marenzelleria neglecta. http://www.nobanis.org/files/factsheets/Marenzelleria_neglecta.pdf

Norkko A; Bonsdorff E; Boström C, 1993. Observations of the polychaete Marenzelleria viridis (Verrill) on a shallow sandy bottom on the South coast of Finland. Memoranda Societatis pro Fauna et Flora Fennica, 69:112-113.

Norkko A; Enberg M; Bonsdorff E, 1995. Occurrence and population dynamics of Marenzelleria viridis. Tvärminne Studies, 6:41.

Olafsson EB, 1986. Density dependence in suspensionfeeding and deposit-feeding populations of the bivalve Macorna balthica: a field experiment. Journal of Animal Ecology, 55:517-526.

Olenin S, 2006. DAISIE Species factsheet. http://www.europe-aliens.org/speciesFactsheet.do?speciesId=53375#

Olenin S; Chubarova S, 1992. Recent introduction of the North American spionid polychaete Marenzelleria viridis (Verrill, 1873) in the coastal areas of the southeastern part of the Baltic area. In: International Symposium - Function of Coastal Ecosystems in Various Geographical Regions, Gdansk University, Poland.

Pelegri SP; Blackburn TH, 1995. Effect of bioturbation by Nereis sp., Mya arenaria and Cerastoderma sp. on nitrification and denitrification in estuarine sediments. Ophelia, 42:289-299.

Segerstrale SG, 1957. Baltic Sea. Memoirs of Geological Society of America, 67(1):751-800.

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Links to Websites

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Aquatic alien species in German inland and coastal watershttp://www.aquatic-aliens.de/species-directory.htm

Contributors

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22/08/09 Original text by:

Merli Pärnoja, Tallinn University, Estonia

Jonne Kotta, Estonian Marine Institute, University of Tartu, M Mäealuse 10a, Tallinn 12618, Estonia

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