Marenzelleria viridis

Invasion History

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

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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.

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Description

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).

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Taxonomy

Ecology

General:

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).

Food:

Phytoplankton; Detritus

Consumers:

Fishes, birds, crabs

Competitors:

Other polychaetes

Trophic Status:

Deposit Feeder

DepFed

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Habitats

General Habitat	Tidal Fresh Marsh	None
General Habitat	Unstructured Bottom	None
General Habitat	Salt-brackish marsh	None
Salinity Range	Limnetic	0-0.5 PSU
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Tidal Range	Subtidal	None
Vertical Habitat	Endobenthic	None
Vertical Habitat	Planktonic	None

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Tolerances and Life History Parameters

Minimum Salinity (‰)	0	Adults frequently occur in fresh water of estuaries (George 1966
Maximum Salinity (‰)	32	Field, Nova Scotia (George 1966; Lab, Netherlands (Schiedek 1999)
Minimum Reproductive Temperature	10	George 1966, worms from Nova Scotia, development incomplete at 2 C
Minimum Reproductive Salinity	10	Larvae were unable to complete development at salinities below 10 ppt (George 1966)
Minimum Duration	24	Larval Period, 20 C- George 1966, worms from Nova Scotia
Maximum Duration	40	Larval Period, 10 C, 30 PSU- George 1966, worms from Nova Scotia
Minimum Length (mm)	40	Dauer et al. 1980
Maximum Length (mm)	140	Nova Scotia (George 1966)
Broad Temperature Range	None	Cold-temperate-Warm temperate
Broad Salinity Range	None	Limnetic-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.

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Regional Impacts

B-II	None		Ecological Impact	Habitat 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-XI	None		Ecological Impact	Competition
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-VII	None		Ecological Impact	Competition
Some community impacts in Vistula Lagoon (Zaiko et al. 2011). Marenzellaria is now dominant in Vistula Lagoon infauna (Ezhova et al. 2005).
B-VII	None		Ecological Impact	Habitat 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-IV	None		Ecological Impact	Competition
A moderate level of community impacts (Zaiko et al. 2011).
B-IV	None		Ecological Impact	Habitat Change
Some habitat impacts (Zaiko et al. 2011).
B-III	None		Ecological Impact	Habitat Change
Some habitat impacts (Zaiko et al. 2011), mostly due to burrowing and bioturbation.
B-X	None		Ecological Impact	Habitat Change
Aeration of sediments, mitigating hypoxia (Norrko et al. 2011)
B-II	None		Ecological Impact	Competition
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-III	None		Ecological Impact	Bioturbation
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).

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