Marenzelleria neglecta -
SCIENTIFIC NAME
Marenzelleria neglecta Sikorski & Bick, 2004
1991 - 2000
NW Atlantic
Shipping (ballast water)
The red-gilled mud worm is native to the Atlantic coast of North America, where it generally occurs in estuaries, coves and bays with brackish water. The red-gilled mud worm in Europe probably originates from the Chesapeake Bay and Currituck Sound in the United States [2].
First observation in Belgium
The red-gilled mud worm was first observed in Belgium on the 23rd of October 1996 in the Sea Scheldt near Doel [3]. Then, it was misidentified as M. viridis. Since using updated identification keys, all individuals from the Western Scheldt got identified as red-gilled mud worms. According to this new key, M. viridis has not been observed in the Scheldt [4]. However, it cannot be excluded that M. neglecta might occur alongside M. viridis. These two species are found together in the Elbe estuary [5, 6].
Spreading in Belgium
The red-gilled mud worm is commonly observed in the Sea Scheldt, even upstream of Antwerp. However, the worm was incompletely identified (or misidentified) as Marenzelleria spp. or Marenzelleria viridis [7, 8]. In our study area, the worm is present in the Dutch Western Scheldt [4] and the Ghent-Terneuzen Canal [9].
Spreading in neighbouring countries
In Europe, the red-gilled mud worm occurred for the first time in 1985 in the Darss-Zingst Bodden Chain, located in the German part of the Baltic Sea. From there, it spread and colonised the entire Baltic Sea [2]. Since the 1980s, the species has been found in the Baltic Sea together with M. viridis and M. arctia [10]. Morphologically, these three species are very similar and, therefore, difficult to identify. An important identification feature is the length of the nuchal organ, which is visible after treatment of the worm with a dye. Numeric features must take size into account
In 1996, the species was found in the German coastal waters of the North Sea, in the Elbe estuary [11], and later in the Weser estuary [2]. The introduction to these estuaries probably occurred via the Kiel Canal, which connects the Baltic Sea with the Elbe estuary [2]. In 2000 – and possibly even in 1993 [9] – the species appeared in the North Sea Canal in the Netherlands [2], and in 2006 in the Limfjord (Denmark) [19]. Since 2014, M. neglecta is found at the mouth of the Don River in the Sea of Azov [13]. The species has not yet been reported in British waters [20].
Most likely, the red-gilled mud worm arrived in Europe from North America via the ballast water of ships. The larvae move freely in the water column and might end up in the ballast water tanks [21]. Once in Europe, this alien species probably continued to spread because of shipping between European ports [22].
The red-gilled mud worm feeds by taking up food particles from the bottom and likely does so more efficiently than some native species, such as the ragworm Hediste diversicolor. In this way, the non-native species can win the competition for food from native species that share the same food sources [23]. This was observed in the Baltic Sea, where it has become a dominant species in several areas. In the Western Scheldt, this is not yet the case and the red-gilled mud worm is currently only sporadically observed [4].
The optimal environment of the red-gilled mud worm is nutrient-rich, brackish water. The species lives in estuaries, lagoons, protected coastal areas and brackish channels with a low to medium salinity (0.5-10 PSU) [8], although it can tolerate varying salinities (0.5-30 PSU) [24]. By comparison, the seawater of the North Sea has a salinity of about 35 PSU.
The red-gilled mud worm can survive at low temperatures but prefers temperatures above 10°C [24, 25]. This alien species is described as one of the best-adapted bristle worms to oxygen-depleted conditions. This polychaete is also resistant to hydrogen sulphide, a highly toxic gas released by the decomposition of organic substances [25].
The red-gilled mud worm has become a dominant species in several habitats of the Baltic Sea. Estimates of maximum densities are extremely high and range from 50,000 [2] to 270,000 individuals per m2 [23]. Such mass populations likely have an impact on the local fauna.
After the introduction of this alien species, populations of native species, such as the ragworm Hediste diversicolor and the amphipod Monoporeia affinis, were negatively affected [23]. Experiments demonstrated that the mortality of H. diversicolor is higher when the red-gilled mud worm is present in the same environment [23]. On the other hand, in the eastern part of the Gulf of Finland, the presence of the worm has not yet affected native species. This can be explained by the low numbers and biomass of this species at this specific location. Furthermore, the low salinity and an alternation of hypoxic and anoxic conditions prevent the further expansion of the species [26, 27].
However, the red-gilled mud worm does not always win. Densities of the red-gilled mud worm are much lower at high concentrations of the hardshell tellin Limecola balthica. The hardshell tellin is not only dependent on the food particles it absorbs from the soil, but – in contrast to the red-gilled mud worm – is a very efficient filter feeder. This makes it easier for the hardshell tellin to cope with food shortage than the non-native species. Since the hardshell tellin is one of the most common species in soft substrates, the food competition between the two species plays an important role in inhibiting the further spread of the red-gilled mud worm [28]. In the Gulf of Finland, competition between the red-gilled mud worm and the hardshell tellin is not observed. The absence of this competition is typical for eutrophic waters with high primary plankton production and low diversity of soil fauna [26, 29-31].
This alien species does not only causes negative effects. For example, the red-gilled mud worm was found in the stomachs of some fish species, indicating that the worm is part of their diet [22, 24]. The species also increases the oxygen concentration in the soil through its digging behaviour – often deeper than many native species. In turn, this digging stimulates other soil organisms and accelerates the decomposition of organic matter [24]. The latter may be a problem in soils where deep-digging species are usually absent
Along the German coast of the Baltic Sea, the vast majority of native species increased in number after the introduction of the red-gilled mud worm. It is currently unclear whether the red-gilled mud worm also affects native species in the Scheldt [7].
To prevent the introduction of these and other species to new places, the International Maritime Organization (IMO) is now calling for ballast water tanks to be cleaned in the open sea so that the organisms present in the tanks are not carried to the final destination [34]. Ballast water may also be treated chemically [21]. These measures have been in force since the 8th of September 2017 [35, 36].
The red-gilled mud worm can grow up to 15.7 cm long and up to 3.2 mm wide [25]. This ground-dwelling creature lives in a mucus-covered J-, L- or I-shaped tube [10, 18, 37-39] that can be up to 25-35 cm deep [3, 22] with an average diameter of 2 mm [10, 18, 37]. It prefers soils rich in silt [3, 22]. Compared to other polychaetes (e.g. Marenzelleria arctia), this species is a relatively ineffective bioturbator. The species only digs dead-end vertical burrows. The absence of a network of burrows and the small tube opening of 1-2 mm reduce the ease of transport of particles in and out of the sediment [40].
The red-gilled mud worm feeds on particles from the soil. Freely suspended microscopic organisms and organic matter can be taken up after being deposited on the soil [25].
The worm reaches sexual maturity after one year [25]. The larvae float freely in the water column for several weeks, after which they begin their life stage in the sediment [17]. The worm has a lifespan of about three years [25].
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