Marenzelleria neglecta

Invasion History

First Non-native North American Tidal Record: 1991
First Non-native West Coast Tidal Record: 1991
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Marenzelleria neglecta is native to the Northwest Atlantic, but its full range is somewhat uncertain, owing to its long confusion with M. viridis. The type specimen was from the Darss-Zingst lagoons on the Baltic coast of Germany, where it is a recent invader. Specimens from Chesapeake Bay; Currituck Sound, North Carolina; and the Ogeechee River estuary, Georgia (Sikorski and Bick 2004; Blank and Bastrop 2009) are presumed to be native. Marenzelleria viridis appears to have a more northern range, from Nova Scotia to Delaware Bay (Sikorski and Bick 2005; Blank and Bastrop 2009). Marenzelleria viridis is believed to penetrate estuaries to a lesser degree then M. neglecta, although adults of both species tolerate freshwater and larvae develop at low salinities (> 5 PSU, M. neglecta, Bochert et al. 1996; Bochert 1997; >10 PSU, M. viridis, George 1966). More sampling of these species on the East Coast of North America is needed, given these worms’ interesting life history, and their ecological importance in their native and invaded ranges.

Polychaetes of the genus Marenzelleria were first collected in the upper reaches of the San Francisco estuary in 1991, and initially identified as M. viridis (Cohen and Carlton 1995). However, subsequent identification also found M. neglecta in the area (Sikorski and Bick 2004; Leslie Harris, personal communication) and more sampling is needed to determine the relative abundance of both species.

In the Baltic and North Seas, three species of Marenzelleria - M. viridis, M. neglecta, and M. arctia - have been introduced in the last three decades, resulting in considerable taxonomic confusion (Bastrop et al. 1995; Sikorski and Bick 2004). The species show some geographical and salinity preferences, with M. viridis occurring mostly in more saline regions of the North Sea coast; M. neglecta in more brackish habitats of the central and southern Baltic, and some low-salinity North Sea estuaries; and M. arctia in the Gulf of Bothnia, but considerable overlap occurs (Bastrop and Blank 2006). The earliest known record of M. neglecta is from 1985 in the Darss-Zingst lagoons, in eastern Germany (Bick and Burckhardt 1989, cited by Sikorski and Bick 2004).

North American Invasion History:

Invasion History on the West Coast:

Marenzelleria spp. was first found in the San Francisco estuary in 1991 at Collinsville, California, on the Sacramento River. It was subsequently found in the Delta at Grizzly Bay, and in wet years, in San Pablo Bay and the uppermost South Bay. It appears to be most abundant and widespread in the estuary after wet years, at limnetic to mesohaline levels (Cohen and Carlton 1995; Peterson and Vayssieres 2010). These worms were also reported from the Napa River in 2004 (Cohen et al. 2005). The worms were initially identified as M. viridis, but Sikorski and Bick (2004) identified 15 specimens from Grizzly Bay as M. neglecta, in their description of this species. Leslie Harris has re-identified some specimens, including those from the Napa River, as M. neglecta (Leslie Harris, personal communication 2014). Environmental conditions in the Delta appear to be closer to the preferences of M. neglecta than those of M. viridis (Bochert 1996; Bochert et al. 1997; Sikorski and Bick 2004). However, an extensive examination of specimens will be needed to determine whether more than one species of Marenzelleria is present in the estuary (Leslie Harris, personal communication 2014).

Invasion History Elsewhere in the World:

Baltic populations of Marenzelleria were first reported in 1985 from the Darss-Zingst lagoons in eastern Germany, on the western Baltic (Bick and Burckhardt 1989, cited by Essink 1999). These were initially assumed to be M. viridis, dispersed from recently discovered populations in North Sea estuaries (discovered in 1979-1986 from Scotland, Netherlands, and Germany; Elliott and Kingston 1987, cited by Bastrop et al. 1998; Atkins et al. 1987; Essink 1999). However, differences in reproductive timing, salinity tolerance of larvae, and genetics led to the suspicion that North Sea and Baltic populations represented at least two separate species (Bastrop et al. 1995; Bastrop et al. 1998; Bastrop and Blank 2006). Subsequent genetic studies revealed that three species were present in European waters, and that their ranges overlapped to some extent. Marenzelleria neglecta was found in at least two North Sea estuaries, the upper Elbe estuary, Germany in 1996 (Sikorski and Bick 2004, 0.7 PSU) and in the North Sea Canal, Netherlands in 2001 (Moorsel et al. 2010, 6-17 PSU). In much of the central Baltic, M. neglecta appears to be the predominant form, but M. viridis occurs at some stations. Marenzelleria neglecta was found with M. arctia at one station in the southern Gulf of Bothnia (Sandviken, Sweden), but at other stations and the Gulf of Finland M. arctia was the only species (Bastrop and Blank 2006; Blank et al. 2008). NB: gaps in the Baltic portion of our distribution map are regions where Marenzelleria spp. were not clearly identified to species by morphological or molecular methods.

In 2008, a spionid identified as M. viridis was collected at one station in the Urias estuary, near Mazatlan, 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. This record could refer to M. neglecta, but further investigation is needed. 

Sikorski and Bick (2004) mention one puzzling record for M. neglecta from Tuktoyaktuk, Northwest Territories, Canada, on the Arctic Ocean. However, specimens were incomplete, so their identity was somewhat uncertain (Sikorski and Bick 2004).

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Description

The body of Marenzelleria neglecta consists of up to 250 chaetigers (chaetae-bearing segments). The prostomium is usually bell-shaped, rounded or bilobed anteriorly, and bears two pairs of eyespots, in a line or with the posterior pair closer together. The palps are short, and do not extend beyond chaetiger 10 in preserved specimens. The nuchal organ reaches the mid-segmental ciliated band of chaetiger 4. Juvenile worms, with 15 chaetigers, have a single pair of branchiae (gills) on the 1st chaetiger, while large worms have up to 69 pairs. The more posterior branchiae tend to diminish in size before disappearing. The parapodia are biramous, with upper notopodia and lower neuropodia each bearing chaetae. On the 1st chaetiger, the lamellae of the notopodia are usually longer than the branchiae, but on subsequent chaetigers the branchiae are up to 1/3 longer. On chaetigers 2-9, the tips of the lamellae are not fused to the branchiae. The upper tips of the notopodial lamellae are usually pointed, but sometimes rounded, and diminish in size posterior, becoming triangular or oval. The neuropodial lamellae are sometimes pointed on anterior chaetigers, but become rounded or triangular posteriorly (Skikorski and Bick 2004). Some of the chaetae take the form of stout hooded hooks on the neuropodia of chaetigers 11 to 51, with 2-8 in each bundle of chaetae. Hooded hooks also occur on the notopodia of chaetigers 12-67, with 2-7 per bundle. The neuropodia have an inferior tuft of chaetae, with bundles of 2-6 chaetae in chaetigers 1-5, with the chaetae becoming fewer and stouter on chaetigers 4-41. The pygidum (anal segment) of juveniles has four pairs of setae, while adults have up to seven cirri. Specimens are up to 115 mm long (Sikorski and Bick 2004). The color of the body varies from green to brown, but the branchiae are always red (Olenin, in DAISIE 2009).

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Taxonomy

Ecology

General:

Marenzelleria neglecta is an estuarine spionid polychaete, typically found in areas of highly variable salinity, but preferring low salinity. Early accounts of its life history as 'M. cf. viridis' are difficult to interpret, 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). Spawning in a Baltic population (probably M. neglecta) occurs in September-December (Bochert 1997). Adults apparently die after spawning, showing semelparous reproductive strategy and investing all available resources into maximizing reproduction (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). Larvae of Baltic M. neglecta settled at 15-23 chaetigers (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-chaetiger 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 upstream 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 M. viridis and M. neglecta inhabit mudflats and shallow muddy bottoms, usually in areas of variable or low salinity (George 1966; Atkins et al. 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 closely related species are apparently similar in feeding and sediment-dwelling behavior, but size and burrowing depth indicate differences in bioturbation and the biogeochemical consequences (Renz and Forster 2013). that can be influenced both by species density and sediment type. 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). Marenzellaria spp. seem to tolerate low oxygen level (Schiedek 1997) and shows characters typical of r- strategists. Due to these it can be classified as opportunistic species (Krauppi et al. 2015). Depth and temperature seem to be the most important variables in accounting for the patterns in Marenzelleria spp. abundance in the Baltic Sea with higher abundances over 10 m depth (Krauppi et al. 2015).

Food:

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	Salt-brackish marsh	None
General Habitat	Canals	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
Tidal Range	Low Intertidal	None
Vertical Habitat	Endobenthic	None
Vertical Habitat	Nektonic	None

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Life History

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Tolerances and Life History Parameters

Minimum Salinity (‰)	0.2	For adult animals, Darst-Zingst lagoon, Germany (Bochert et al. 1997).
Maximum Salinity (‰)	30	Highest tested (Bochert et al. 1997). Fritsche (1997) found a 72-h LC50 of 29.8 for animals acclimated at 5 PSU and 10 C. In San Francisco Bay, the seaward limit in a survey of muddy sediments was 16.1 PSU (Lee et al. 2003).
Minimum Dissolved Oxygen (mg/l)	1	Marenzelleria spp. are characterised by a very efficient use of oxygen during hypoxia, since anaerobic energy production started at a very low oxygen pressure (critical pO, at about 2 kPa) (Schiedek 1997). 2kPA; ~ 1mg/L
Minimum Reproductive Salinity	5	Mimimum for successful maturation of eggs and sperm, and larval development (Bochert 1997)
Maximum Reproductive Salinity	10	Maximum for successful maturation of eggs and sperm, and larval development. However, the next highest salinity tested was 25 PSU (Bochert 1997)
Minimum Duration	30	Egg to settling at 20 C,10 PSU (Bochert et al. 1996)
Maximum Duration	85	Egg to settling at 10 C, 20 PSU (Bochert et al. 1996)
Minimum Length (mm)	30	Animals below 20 mm are all juvenile. Animals >40 mm are adults which have spawned once. (Bochert 1997)
Maximum Length (mm)	115	Sikorski and Bick 2004
Broad Temperature Range	None	Cold temperate-Warm temperate
Broad Salinity Range	None	Limnetic-Polyhaline

General Impacts

The spionid polychaetes Marenzelleria neglecta 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. neglecta, together with M. viridis 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 be 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 promoting P retention and nitrification in the longer term (Hietanen et al. 2007; Hedman et al. 2011; Norkko et al. 2011). Experimental studies with worms and sediments from Odense Fjord, Denmark, showed that introduced M. viridis (which burrows deeper) 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.  A review of polychaete impacts indicates that the three major Marenzellaria (arctia, neglecta, viridis) species may differ in their burrow stucture and impacts, but are often confused and lumped.  As a group, they have the higherts number of publucations on thoioer impacts (Alvarez-Aguilar et al. 2022).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Habitat Change
Mucus from the tubes of Marenzellaria sp. as well as the mucus produced by other native and introduced deposit feeding and tube-building benthos, contributes to a surface layer of flocculent fluff, which may trap much more phytoplankton than that actually consumed by the animals (Jones et al. 2009).
P090	San Francisco Bay		Ecological Impact	Habitat Change
Mucus from the tubes of Marenzellaria sp. as well as the mucus produced by other native and introduced deposit feeding and tube-building benthos, contributes to a surface layer of flocculent fluff, which may trap much more phytoplankton than that actually consumed by the animals (Jones et al. 2009).
B-VI	None		Ecological Impact	Bioturbation
Burrowing activitity of M. neglecta results in reworking of sediment particles and increased irrgiation of sediments, increasing oxygenation and the exchange of solutes. The higher density of worms and higher rates of irrigation suggest that the impact of M. neglecta is greater than that of the native polychaete Hediste diversicolor (Hedman et al. 2011).
B-X	None		Ecological Impact	Toxic
Bioturbation by Marenzelleria neglecta resulted in increased release of toxic organic compounds from sediments in the Stockholm archipelago(Granberg et al. 2008).
B-VIII	None		Ecological Impact	Competition
In experiments, Marenzelleria neglecta reduced the survival of Hediste diversicolor (Kotta et al. 2006).
B-VIII	None		Ecological Impact	Habitat Change
In experiments, Marenzelleria neglecta enhanced the chlorophyll content of sediments, presumably by releasing nuritents, while burrowing (Kotta et al. 2006).
B-IV	None		Ecological Impact	Bioturbation
Deep burrowing by Marenzelleria neglecta results in increased fluxes of nitrate and phosphate from sediments (Renz and Forster 2013; Renz and Forster 2014). However, in experiments, M. neglecta had lower rates of irrigation and survival in high salinity Baltic water and sediment (22 PSU) and low salinity water and sediment (6 PSU) (Quintana et al. 2018).
B-X	None		Ecological Impact	Bioturbation
Irrigation of sediments by Marenzelleria neglecta may result in the oxygenation of sediments, enhancing retention of phosporus in sediments, while reducing the concentration of labile orgnanc compounds (Norkko et al. 2012).
B-X	None		Ecological Impact	Habitat Change
Irrigation of sediments by Marenzelleria neglecta has led to an increase in oxygen levels of sediments, increasing phosphorus retention in sediments, and reducing hypoxia due to eutrophication in the Stockholm archipelago (Norkko et al. 2012)
CA	California		Ecological Impact	Habitat Change
Mucus from the tubes of Marenzellaria sp. as well as the mucus produced by other native and introduced deposit feeding and tube-building benthos, contributes to a surface layer of flocculent fluff, which may trap much more phytoplankton than that actually consumed by the animals (Jones et al. 2009)., Mucus from the tubes of Marenzellaria sp. as well as the mucus produced by other native and introduced deposit feeding and tube-building benthos, contributes to a surface layer of flocculent fluff, which may trap much more phytoplankton than that actually consumed by the animals (Jones et al. 2009).

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