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Aquatic Invasions (2014) Volume 9, Issue 4: in press doi: http://dx.doi.org/10.3391/ai.2014.9.4.01

© 2014 The Author(s). Journal compilation © 2014 REABIC

Open Access

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Supplementary material

Impacts of invasive alien marine species on ecosystem services and biodiversity: a pan-European review

Stelios Katsanevakis1*, Inger Wallentinus2, Argyro Zenetos3, Erkki Leppäkoski4, Melih Ertan Çinar5, Bayram Oztürk6, Michal Grabowski7, Daniel Golani8 and Ana Cristina Cardoso1

1European Commission, Joint Research Centre (JRC), Institute for Environment and Sustainability (IES), Ispra, Italy 2Department of Biological and Environmental Sciences, University of Gothenburg, Sweden 3Institute of Marine Biological Resources and Inland Waters, Hellenic Centre for Marine Research, Ag. Kosmas, Greece 4Department of Biosciences, Environmental and Marine Biology, Åbo Akademi University, Turku, Finland 5Ege University, Faculty of Fisheries, Department of Hydrobiology, Bornova, Izmir, Turkey 6Faculty of Fisheries, Marine Biology Laboratory, University of Istanbul, Istanbul, Turkey 7Department of Invertebrate Zoology & Hydrobiology, University of Lodz, Poland 8Department of Ecology, Evolution and Behavior and the National Natural History Collections, The Hebrew University of Jerusalem, Israel

E-mail: [email protected] (SK), [email protected] (IW), [email protected] (AZ), [email protected] (EL), [email protected] (MEC), [email protected] (BO), [email protected] (MG), [email protected] (DG), [email protected] (ACC)

*Corresponding author

Received: 8 January 2014 / Accepted: 6 June 2014 / Published online: 4 August 2014

Handling editor: Vadim Panov

Online article is available at http://www.aquaticinvasions.net/2014/ACCEPTED/AI_2014_Katsanevakis_etal_correctedproof.pdf

Supplementary material

Supplement 1. Species-specific review of the impacts of invasive alien species on ecosystem services and biodiversity in the European Seas, and other related information. A superscript E or N indicates publications documenting impacts based on manipulative or natural experiments respectively.

Recommended Citation of this material: Katsanevakis S, Wallentinus I, Zenetos A, Leppäkoski E, Çinar ME, Oztürk B, Grabowski M, Golani D, Cardoso AC (2014) Impacts of invasive alien marine species on ecosystem services and biodiversity: a pan-European review. Aquatic Invasions 9 (in press)

Dinophyta (Myzozoa)

Alexandrium minutum: This dinoflagellate (type locality in Alexandria, Egypt) produces various paralytic shellfish toxins, which accumulate in shellfish and can be transmitted through the food chain, as well as to humans. Also non-toxic strains exist e.g. in Italy, on the Scottish east coast and Orkney (e.g. Nascimento et al. 2005 and references therein; Brown et al. 2010; Anglès et al. 2012). The toxicity may also vary depending on environmental conditions (Touzet et al. 2008). In northern Europe, persistent blooms of A. minutum have occurred since 1985 and have caused severe losses to aquaculture (Nehring 1998). Biogeographical analyses using molecular techniques have shown that there is a global clade found in Europe, Australia and South Africa, while populations from New Zealand belong to the Pacific Clade (McCauley et al. 2009). The importance of encystment and excystment of the resting cysts, acting as seed banks to make the blooms recurrent, has been emphasized (Bravo et al. 2010; Anglès et al. 2012). Blooms of this species often occur in nutrient-rich enclosed areas of harbours, lagoons and estuaries, mainly under stable water column conditions. Van Lenning et al. (2007) measured environmental variables and variation in cell numbers during a bloom until it finally declined, which they attributed to grazing. On the other hand, other studies have shown that parasites may be involved in reducing a bloom. In northern Brittany, France, Chambouvet et al. (2008) showed that up to 40% of the cells were infected by species-specific parasites from the dinoflagellate group Syndiniales, believing them to control

S. Katsanevakis et al.

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the blooms of this species. However, Spanish scientists (Figueroa et al. 2008a, 2010), studying blooms from the NW Mediterranean, showed that the recently described perkinsoid parasite Parvilucifera sinerae did not infect the resting cysts, which act as seed banks, but only the mobile planozygots and temporary cysts, and that clones can become resistant by crosses and recombinations. They thus emphasized that there is a need to re-evaluate the use of parasites for controlling toxic blooms. Furthermore, small-scale turbulence has been shown to reduce and delay parasite infections but cannot prevent them (Llaveria et al. 2010). Large blooms consume nutrients, decrease light, which also affects the habitat negatively, and toxic blooms have a negative impact on recreation. Experiments have been conducted to increase the biomass of A. minutum and lipid production in bioreactors, with the goal of producing biofuels (e.g. Fuentes-Grünewald et al. 2013).

Alexandrium monilatum: This chain-forming dinoflagellate, type locality in Indian River, Florida, produces the lipophilic ichtyotoxic goniodomin A (Hsia et al. 2006) causing numerous fish kills and widespread discolouration in warmer waters. It also affects shellfish behaviour and increases larval mortality (May et al. 2010). In Europe, however, it has only been listed from the W Black Sea (Moncheva et al. 2001; Zaitsev and Öztürk 2001; Gomoiu et al. 2002), where it was reported to reach bloom-forming densities with ecological and socioeconomic implications. Large blooms consume nutrients, decrease light, which also affects the habitat negatively, and toxic blooms have negative impact on recreation. However, Gómez and Boicenco (2004) and Gómez (2008) questioned the presence of this species in the Black Sea.

Karenia mikimotoi: This dinoflagellate, type locality in Japan, has been observed to build up large blooms along many European Atlantic coasts (initially misidentified as Gyrodinium aureolum), which have killed both wild and farmed fish, shellfish, and benthic invertebrates such as lugworms, cockles, brittle stars etc. from 1968 onwards (Ottway et al. 1979; Boalch 1987 and references therein; Raine et al. 2001 and references therein; Mitchell and Rodger 2007; Davidson et al. 2009a, 2009b; Rodger et al. 2011; Guiry and Guiry 2013). Chinese scientists showed in experiments and by analyses that the kills were caused by haemolytic substances (2 liposaccharides and 1 lipid), but that also allelochemicals of other compositions had a haemolytic activity (Yang et al. 2011). Air quality is affected by the fact that anoxia is caused by the dying, sinking bloom and by the smell of killed organisms. Large blooms also consume a large quantity of nutrients. The negative effect on recreation can be exemplified by the temporary closure, in 2012, of several beaches in Co. Donegal, which had been affected by a bloom of Karenia mikomotoi, by the Environmental Protection Agency, Ireland (2013). Although the microbiological quality of bathing waters was not affected, large numbers of dead fish and crustaceans had been found in the area. Release of allelopathic chemicals affects other plankton species, and the presence of other dinoflagellates could send water-borne cues, which increased toxin production (Yang et al. 2011). The kills also affect many parts of the ecosystem. The released chemicals decreased light also affecting the habitat negatively. It was shown by molecular tools that isolates from Europe and New Zealand are more closely related to each other than either group is to isolates from Japan (Al-Kandari et al. 2011).

Gymnodinium catenatum: This dinoflagellate, type locality in Mexico, is a chain-forming dinoflagellate that produces gonyautoxins and saxitoxins (Gárate-Lizárraga et al. 2004), metabolites responsible for Paralytic Shellfish Poisoning (PSP). Humans eating contaminated shellfish or fish (i.e. bivalves that filter algal cells from the water and accumulate toxins or fish that have accumulated the toxins through foodwebs) show paralytical disorders and may die because of PSP. The species is responsible for major worldwide losses in aquaculture due to shellfish toxicity. In 1976 there was a major PSP event in the Galician Rías (NW Spain), where nearly 200 cases of PSP occurred (Luthy 1979). Galician Rías Baixas is the major producer of mussels in Europe, and the impact of closures in shellfish aquaculture facilities because of G. catenatum blooms was particularly severe (Ribeiro et al. 2012). G. catenatum has become an abundant and well-established species in the Alborán Sea (westernmost part of the Mediterranean Sea) and is associated with frequent toxic events (e.g., Taleb et al. 2001; Calbet et al. 2002). The toxins produced by G. catenatum do not only accumulate in molluscs, but experiments in other areas, have shown that they also affect their behaviour (e.g. Estrada et al. 2010E; Escobedo-Lozano et al. 2012E). During blooms along the Iberian peninsula these toxins have also been analyzed in various fish species such as the planktivorous sardines (Costa et al. 2010), the horse mackerel, a predator on zooplankton, which itself is eaten by top predators (Lage and Costa 2013), thus contributing to a negative impact on the whole ecosystem, and the seabream (Costa et al. 2012). Also, for cephalopods from the Portuguese coast, analyses showed that their digestive glands contained toxins (Costa et al. 2009), which also contributes to negative food-web effects. Calbet et al. (2002) suggested that the

Impacts of invasive marine species – Supplementary material

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presence of Gymnodinium catenatum in the Alborán Sea has an important impact on mesozooplankton grazing rates and survival, even when they are not at ‘bloom’ concentrations, and thus might have resulted in changes in the pelagic food web. Experiments with aquacultured shrimps in Mexico resulted in mortality and abnormal behaviour at high algal densities, while the shrimps had good survival at low densities (Pérez Linares et al. 2009E) and they emphasized that cultured shrimps represent an unexpected potential risk for humans. During blooms of G. catenatum in Portugal, Costa et al. (2010) were able to measure extracellular toxins in the seawater. Dense blooms reduce the nutrient concentrations for other species and have a negative impact on habitats by reducing light; they also have a negative impact on recreation.

Haptophyta

Phaeocystis pouchetii: This species, first described from northern Norway, can form large colonies originally formed by small free-swimming solitary cells, which become embedded in a matrix of heteropolysaccharidic chains (Schoemann et al. 2005; Veldhuis and Wassmann 2005). Dense blooms reduce nutrients and compete with other phytoplanktonic species; declining colonies produce foam and cause anoxia, which is negative for recreation (Cadée and Hegeman 1986; Riegman et al. 1992; Schoemann et al. 2005 and references therein). Dense blooms can act as ecosystem engineers (sensu Berke 2010) by reducing light for benthic autotrophs. It is a partly native species in Europe occurring along several European coasts and most effects described below are from areas where the species is indigenous. However, such impact may also occur in areas where it is alien. It has been introduced by ballast water discharges into the NW and NE waters of the Black Sea where it can build up blooms (Yasakova 2013 and references therein), while it is not clear if the records from the central Aegean Sea (Moncheva et al. 2001; Ignatiades and Gotsis-Skretas 2010) are introduced or native populations. Colony-forming blooms from near Tromsö, northern Norway, where the species is indigenous, were analysed for toxic compounds (Hansen et al. 2004 and references therein) and effects were found on dividing sea urchin embryos caused by a polyunsaturated aldehyde (PUA). This species has been reported to be grazer deterrent to copepods (Estep et al. 1990E; Hamm 2000), to reduce growth in farmed salmon (Eilertsen and Raa 1995N), and water from the algal cultures was toxic to cod larvae (Aanesen et al. 1998E). This substance may also have caused the long known avoidance of these blooms by herrings. Phaeocystis species are known to produce dimethylsulphoniopropionate (DMSP), which could be due to an excess of energy (Stefels 2000) that can be enzymatically converted into dimethylsulphide (DMS), which is involved in the biological regulation of climate. DMS is a major source of cloud-condensation nuclei (CCN), which regulate the reflectance (albedo) of clouds (Bates et al. 1987; Charlson et al. 1987; Carslaw et al. 2010). DMS emission seems to have a cooling tendency (as CO2 pumping) through its effect on planetary albedo, and thus these DMSP-producers are considered to have a positive impact on climate regulation (but negative on air quality). Schoemann et al. (2005) estimated that the contribution of Phaeocystis blooms to the global DMS flux to the atmosphere is 5 to 10%. In native areas diatoms and other protists have been found on large colonies, which act as a new substrate (Sazhin et al. 2007). It was previously assumed that sinking organic material from Phaeocystis blooms constituted an efficient part of the biological carbon pump (e.g. Hamm 2000), but later estimates by Reigstad and Wassmann (2007) from northern Norway showed that they only contributed around 3% of the vertical POC flux at 100 m depth and <5% of that was due to the mucus.

Ochrophyta

Coscinodiscus wailesii (initially misidentified as Coscinodiscus nobilis; should not be considered as alien according to Gómez and Souissi 2010): This very large diatom, first described from Pudget Sound, NE Pacific, was first recognized, in Europe, in the English Channel in winter-spring 1977, when a bloom was observed producing enormous amounts of slime, which clogged trawls and by accumulating clay particles made fishing difficult (Boalch and Harbour 1977 as C. nobilis; Boalch 1984, 1987; Edwards et al. 2001). It then sunk to the bottom accumulating in certain areas and large amounts were swept into the trawls. Also, in culture, it produced a lot of slime, the measured volumes per cell being used to estimate


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the slime production in sea. However, the slime does not always seem to be produced (Boalch 1987), and it has not been possible to find any recent studies confirming the same drastic impact. In Japan, where it is a native species, bleaching of farmed nori has been reported, caused by less nutrients being available for the nori during the bloom of this algal species, which often starts from resting cells in the sediment (Nishikawa et al. 2010). Field data from the German Bight and laboratory experiments showed that this species accumulated 8–10 times less copper and 10–20 times less cadmium and zinc than other abundant phytoplankton species, suggesting it has a higher tolerance to higher doses of heavy metals in inshore regions than the native species (Rick and Dürselen 1995). They also measured that during a bloom it could assimilate up to 90% (up to 1.4 mg C l-1) of phytoplankton carbon and since many, but not all grazers avoid it because of its huge size (Roy et al. 1989; Jansen 2008), this can mean a reduced transfer of carbon within the food web (Rick and Dürselen 1995). On the other hand, decapods such as the prawn Crangon crangon feed well on C. wailesii and thus may benefit from the mass occurrence of the diatoms (Nehring 1999). The huge slime production, especially when mixed with clay and dead organisms, may have a negative effect on recreation. Substantial damage is caused if the copious mucilage sinks and covers the seabed, likely causing anoxic conditions (DAISIE 2013). On the other hand, its huge cells have been used in many basic experiments in ecology, morphology and physiology (e.g. Scheffel et al. 2011; Rzadkowolski and Thornton 2012).

Fibrocapsa japonica: This raphidophycean alga, first described from Atsumi Bay, SE Japan, is potentially toxic. Laboratory experiments have shown that sole larvae died when directly exposed to cells of this species being in late exponential growth phase (de Boer et al. 2012E). These authors also referred to other studies in which other species of fish have been affected. After long term exposure to high cell densities, Pezzolesi et al. (2010)E showed that this species also can cause death to crustaceans and sea bass. In The Netherlands, dense blooms occur yearly (de Boer 2006 in de Boer et al. 2012), which can have negative impacts on recreation and light conditions. Allelochemical compounds can be released, which have been shown to inhibit bacterial luminescence and through bioassays have been used to quantify the negative effects on bacteria and may result in more favourable growth conditions for F. japonica (van Rijssel et al. 2008E).

Odontella sinensis: For this cryptogenic diatom, information about its impact on ecosystem services and biodiversity is insufficient.

Pseudochattonella verruculosa: This dictyochophycean alga, first described from the Seto Island Sea (west of Osaka, Japan), is potentially toxic, and can kill fish. The toxin is probably a fatty acid. The gill tissue of fish is also affected. Laboratory bioassays were run to study the toxic impact of two strains of this species on fish, as well as nine strains of the related native species P. farcimen. The experiments showed that metabolites in P. verruculosa affected the fish cell lines and gills of cod and salmon larvae; stronger effects were found by the strain isolated in Japan than the one from the North Sea (Skjelbred et al. 2011E). Dense blooms have been observed during several years in northern Europe (for references see Skjelbred et al. 2013), which negatively affect nutrient concentrations and light, as well as recreation. P. verruculosa caused the death of 350 tonnes of farmed Norwegian salmon in 1998 and 1100 tons in 2001 (Edvardsen et al 2007). There were also reports on mortality among wild fish (garfish, herring, sandeel and mackerel) along the west coast of Denmark (Aure et al. 2001; Bourdelais et al. 2002; Naustvoll 2010). Another bloom in 2001 also resulted in economical loss for the affected salmon farms (Naustvoll 2010). Apart from its native region in Japan, these phenomena, including fish mortality, have also been recorded from New Zealand (MacKenzie et al. 2011).

Macroalgae

Acrothamnion preissii [Rhodophyta]: This Australian red alga was detected in 1969 on the coast of Tuscany, Italy, NW Mediterranean Sea, most likely having arrived as fouling on ship hulls (Boudouresque and Verlaque 2002), and later spread. Today these filamentous, branched algae form dense turfs in many areas in the western basin on the matte and rhizoms (often dominant on the latter) of the seagrass Posidonia oceanica (e.g. Piazzi et al. 1996; Piazzi and Cinelli 2001, 2003N). It reduces the diversity of these seagrass meadows and can also clog fishing gear (Cinelli et al. 1984) and thus has a negative impact on coastal fisheries. On rocks it is now often replaced (Piazzi and Cinelli 2001, 2003) by a later introduced red alga, Womersleyella setacea (see below), but on other rocky shores it is, even


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without W. setacea, only found in low abundances (Sales and Ballesteros 2010). It has also been found growing over calcareous red algae in maerl beds in the Baleares (Ferrer et al. 1994), where accumulation of sediments may harm these coralline algae. The dense turfs become efficient sediment traps, a positive impact on coastal protection (Piazzi et al. 1996), although the Posidonia beds themselves are quantitatively more efficient in that process (Ballesteros et al. 2007). Dense turf of these thin filamentous plants have a high surface to volume ratio, resulting in high uptake rates of nutrients and carbon dioxide (Littler 1980; Littler and Littler 1980; Wallentinus 1984), meaning a negative impact on the availability of nutrients for coarser algae. The trapping of sediments has a negative impact on the aesthetic value of the ecosystem as well as on recreational activities, such as snorkelling and SCUBA diving. The same negative impact applies to the life cycle maintenance of other species, partly due to the reduction in diversity, but also to the trapping of sediments. The sediment prevents other species, relying on the settlement of spores, to find a solid substrate for settlement, while A. preissii itself can rely on vegetative fragments, which can grow into new plants (e.g. Piazzi and Cinelli 2000N). Their superior performance versus other algae means a negative impact on many species. Ecosystem services provided by communities of sublittoral algae degraded due to A. preissii, such as food, biotic materials, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are thus negatively impacted. Reduced recruitment and increased mortality of gorgonian juveniles have been reported because of A. preissii and other turf algae (Linares et al. 2012E). Since it also grows on the leaves of the seagrass P. oceanica, albeit to a lesser degree as regards bottom coverage, it may have a negative shading effect on this keystone species. This dominant turf species is avoided by the key herbivore of Posidonia-meadows, the sea urchin Paracentrotus lividus (Tomas et al. 2011bE). It also acts as a habitat/ecosystem engineer (Wallentinus and Nyberg 2007; Berke 2010) by negatively affecting the available substrate for other epiphytic macroorganisms to settle on, and the dense turf also reduces the light for other species. On the other hand, it may provide more surface area for microorganisms to grow on than the seagrass plants do. It has not been possible to find any cognitive benefits for this species.

Asparagopsis armata [Rhodophyta]: This red alga was originally described from Western Australia, and was first recorded in Europe from western France in 1925. It belongs to the same family as Bonnemaisonia hamifera and has a heteromorphic life cycle with small, tufted, tetrasporophytes (often referred to as the Falkenbergia rufolanosa phase) and branched gametophytes being some 10s of cm high. The hooked gametophytes have a negative impact on coastal fisheries by clogging fishing nets (Streftaris and Zenetos 2006) and can also grow on oysters (Mineur et al. 2007). Farming of vegetatively propagated gametophytes growing on ropes started in Brittany, NW France, in the mid 1990s to extract bioactive substances (mainly brominated and iodinated methanes and acetones) to be used in the cosmetic industry and in medical products; such farming has also been developed in Ireland since the end of the 1990s (Kraan and Barrington 2005). Schuenhoff et al. (2006)E suggested the use of A. armata as a biofilter in aquaculture. It has positive effects on water purification and Mata et al. (2010)E showed that tanks with tetrasporophytes of this species were more efficient as a biofilter for effluents from a fish farm in Portugal than the conventionally used Ulva rigida. Growing in tanks with fish pond effluents in S Portugal, Figueroa et al. (2008b)E also showed their efficiency as a biofilter and pointed to their potential as a source for mycosporine-like amino acids as UV screen photoprotectors. The tetrasporophytes, by their high ratio of surface to volume, have a greater potential for rapid uptake of nutrients in comparison to their host algae (Littler 1980; Littler and Littler 1980; Wallentinus 1984), a negative impact on ocean nourishment. When the tetrasporophytes cover their host algae densely, less light and nutrients reach them. Extracts of A. armata were shown by Paul et al. (2006a) to have antibacterial activity against marine (Vibrio spp.) and biomedical (Escherichia coli, Pseudomonas aeruginosa and Staphylococcus spp.) strains. Furthermore, these compounds, bromoform and dibromoacetic acid, also reduced the densities of epiphytic bacteria growing on normal A. armata in comparison to on plants for which bromine had not been available during culture. Using the same kind of bromine-deficient tetrasporophytes Paul et al. (2006b) also showed that two natural Australian mesograzers (an amphipod and young abalone) avoided plants cultured with bromide, while the native sea hare Aplysia parvula was not deterred. The same species of Aplysia was also shown to be able to detect the least toxic stage of A. armata, the male gametophytes, which were also the most nutrient-rich, resulting in fewer female plants by the end of the growing season (Vergés et al. 2008). In the NW Mediterranean areas with high densities of grazing fish, Sala and Boudouresque (1997) showed that most of the native fleshy algae had been grazed away. In those areas, the chemically defended A. armata


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gametophytes, being competitively inferior, were totally dominant during late spring. Later, in early summer, large amounts of dead plants accumulated in deeper pools, having a negative effect on recreation. Medical tests were carried out by Zubia et al. (2009) on human cancer cells, to elucidate if extracts of this species and other French red algae had any cytotoxic activity, which they found in A. armata. Furthermore, extracts from this and the closely related A. taxiformis were found by Genovese et al. (2009) to have an antiprotozoan activity, which has the potential to develop into products that could stop the serious parasitic human disease Leishmaniasis. Dichloromethane extracts were obtained from A. armata, the only of the 26 tested seaweeds that exhibited a strong activity against all five fish pathogenic bacteria that Bansemir et al. (2006) included, and thus have the potential to be an alternative to conventional antibiotics.

Asparagopsis taxiformis [Rhodophyta]: This red alga, also belonging to the Bonnemaisoniaceae, was first described in 1813 from Alexandria, Egypt, but the species has a wide distribution in tropical and subtropical areas. It has a heteromorphic life cycle with small, tufted tetrasporophytes (often referred to as the Falkenbergia hillenbrandii phase) morphologically quite difficult to separate from the tetrasporophytes of A. armata [but see characters on morphology and ecophysiological performance given by Ní Chualáin et al. (2004)]. For the occurrence in the Mediterranean Sea, Zenetos et al. (2010) listed one group of this species as being of possible Atlantic origin or cryptogenic, and one invasive strain as being Indo-Pacific. A more detailed explanation was given by Andreakis et al. (2009) and references therein, who provided evidence that the alien invasive, polyploid strain, also found in the Pacific, was first recorded from Sicily in 2000, and then in a few years spread throughout the western basin and to the Atlantic south coast of Portugal. A. taxiformis has a potential to be farmed for the production of economically valuable secondary metabolites (bromoform and dibromoacetic acid) (Mata et al. 2011, 2012). Since the tetrasporophytes are morphologically very difficult to distinguish from A. armata, they probably have the same capacity as biofilters as those of the closely related species (cf. Mata et al. 2010). The tetrasporophytes, due to their high ratio of surface to volume, have a greater potential for rapid uptake of nutrients in comparison to their host algae (Littler 1980; Littler and Littler 1980; Wallentinus 1984), a negative impact on ocean nourishment. Ethanol extracts of a strain from the Strait of Messina were screened for antibacterial activity against pathogenic shellfish and fish bacteria previously isolated from local marine and brackish environments (Genovese et al. 2012). Their results indicated that these extracts could represent a source of antibacterial substances with a potential for use in aquaculture. Furthermore, Australian experiments (Hutson et al. 2012) showed that water-soluble extracts from A. taxiformis could be used to reduce the hatching success of a monogenean ectoparasite, for which there are no methods for preventing infections, and to prolong the time between first and last hatch of the parasites among farmed barramundi. Extracts from this species and the closely related A. armata were found by Genovese et al. (2009) to have an antiprotozoan activity, which has the potential to develop into products that could stop the serious parasitic human disease Leishmaniasis. Analyzing A. taxiformis Jha et al. (2013) showed for the first time for this species that, besides antibacterial activity, it also had a strong antiquorum, sensing activity, which they saw as a strong potential to be used in antifouling compounds against marine biofilms. Considering that there are two different mitochondrial lineages of this species in the Mediterranean it is not surprising that the reports on their impact differ. Papers from the eastern basin often report low abundances with no invasive behaviour (e.g. Tsiamis and Panayotidis 2007), although since then it has exhibited rapid spread, especially in the South Aegean Sea (Tsiamis et al. 2010). In the westernmost Mediterranean (the Alborán Sea), where A. taxiformis was first recorded in 1999, Altamirano et al. (2008) reported that it was rapidly spreading to the Andalusian coast of southern Spain, after it was first introduced in the late 1990s and, in some areas, the gametophytes formed conspicuous monospecific stands and appeared to act as a keystone species. The dispersal was explained by vegetative reproduction. In some areas, it was observed growing on the rhizomes of the seagrass Posidonia oceanica, forming self-sustained stands in protected areas, and has invaded areas previously occupied by A. armata. On the NW coast of Italy, Tamburello et al. (2013) showed that A. taxiformis dominates assemblages in topographically complex habitats, where it coexists with an extensive cover of Caulerpa cylindracea, while A. taxiformis is virtually absent on homogenous platforms, even in experiments where other algae were removed, depending on heterogeneity in environmental conditions.

Bonnemaisonia hamifera [Rhodophyta]: This red alga from the NW Pacific, with a heteromorphic life cycle, was first recorded in Europe as early as 1890 (as tetrasporophytes – previously called


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Trailliella intricata), probably having arrived by shipping, while in the Mediterranean it only dates back forty years. Nash et al. (2005) performed experiments, which showed that this species has a great potential to be farmed for producing secondary metabolites that can be used as preservatives in industrial applications (cf. Asparagopsis armata); however, to our knowledge, the species is not yet used industrially for the provision of such materials. In experiments, Nylund et al. (2013) showed that there was a high cost for producing the secondary metabolites, functioning as grazer deterrents and excreted allelopathic compounds (see references below). The species’ tetrasporophytes, by their high ratio of surface to volume, have a greater potential for rapid uptake of nutrients in comparison to their host algae (Littler 1980; Littler and Littler 1980; Wallentinus 1984), which is perceived as a negative impact on ocean nourishment. Medical research has shown that ethanol extracts of B. hamifera can scavenge UVB radiation-induced reactive oxygen species, which reduces the risk of damage to human keratinocytes (Piao et al. 2012a). Furthermore, experiments in basic ecological theory with this species have shown that the grazer deterrent 1,1,3,3-tetrabromo-2-heptanone, also works as an allelopathic compound. For the first time, Svensson et al. (2013)E showed that this compound can be transferred from this alien alga to a native host algal species by direct contact, with an active and unaltered function, i.e. in inhibiting recruitment of native competitors. Experiments by Enge et al. (2013)E showed that native herbivores strongly preferred native algae to B. hamifera, which was tracked down to the algal metabolite 1,1,3,3-tetrabromo-2-heptanone, not known from the native algae in the invaded area, but also that herbivores used B. hamifera as a refuge for fish predation. The importance of the chemical defence was further underlined by the feeding preference of herbivores for B. hamifera individuals with an experimentally depleted content of this substance (Enge et al. 2012E), the first conclusive example of a highly successful invader, where low grazing pressure in the new range can be directly attributed to a specific chemical defence. The same secondary metabolite was investigated (Persson et al. 2011E) by using field panels coated with the metabolite at concentrations covering those measured at the algal surface. This significantly affected the natural fouling community by altering the composition, and changed the diversity by increasing the evenness and decreasing the density, indicating a broad specificity of this metabolite against bacterial colonization. B. hamifera has a negative impact as an ecosystem engineer by forming dense epiphytic growth on host algae (e.g. Johansson et al. 1998), thus reducing light and nutrient availability for those algae, in addition to preventing competing algae to colonize (Svensson et al. 2013E).

Caulerpa cylindracea [Chlorophyta]: Until recently, the species was accepted as Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman and Boudouresque, but it is now reinstated to its species rank as C. cylindracea Sonder (Belton et al. 2014). The species may form compact multilayered mats up to 15 cm thick that trap sediment; an anoxic layer may develop underneath (Piazzi et al. 2007N; Klein and Verlaque 2008). Its stolons can quickly elongate and easily overgrow other macroalgal (Piazzi et al. 2001N, 2005E; Piazzi and Ceccherelli 2006E) or invertebrate species (Kružić et al. 2008; Baldacconi and Corriero 2009N; Žuljević et al. 2010) causing their elimination. Important changes in the biocommunities of the invaded areas have been reported (Argyrou et al. 1999N; Piazzi et al. 2001N, 2005E, 2007N; Piazzi and Cinelli 2003N; Balata et al. 2004N; Piazzi and Ceccherelli 2006E; Klein and Verlaque 2008, 2011E; Bulleri et al. 2010E). Both negative and positive impacts of C. cylindracea on the seagrasses Cymodocea nodosa and Zostera noltii have been reported (Ceccherelli and Campo 2002E): shoot density of Cymodocea nodosa decreased in the presence of C. cylindracea whereas it increased for Zostera noltii; in both seagrasses the frequency of flowering shoots increased. C. cylindracea affects the performance of Posidonia oceanica causing reduced leaf length and leaf area index and an increase in primary foliar production and in the number of leaves produced annually (Dumay et al. 2002N). Several biotopes, such as Mediterranean communities of sublittoral algae and coralligenous communities, are affected by C. cylindracea, which smothers indigenous populations, outcompetes native communities, and diminishes the structural complexity and species richness (Klein and Verlaque 2008; Katsanevakis et al. 2010; Salomidi et al. 2012). Hence, ecosystem services provided by these biotopes, such as food, biotic materials, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are negatively impacted. The large-scale modification of submarine landscapes induced by C. cylindracea with the overgrowth of benthic assemblages by a more or less dense and continuous web-like green meadow may reduce the attractiveness of the biota for underwater tourism (Klein and Verlaque 2008). Mediterranean seagrass meadows, particularly those of the endemic species Posidonia oceanica, are experiencing widespread decline due to a combination of direct anthropogenic


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pressure and climate change; zones of fibrous remnants of former Posidonia oceanica beds (‘matte morte’) are very vulnerable to C. cylindracea invasion (Katsanevakis et al. 2010), which, however, protects sediments from erosion as well as or even better than seagrasses (Hendriks et al. 2010E). On previously unvegetated soft bottoms, C. cylindracea may be considered as an ‘ecosystem engineer’ building up different microhabitats and shelters, providing new sources of food, promoting the establishment of many zoo-benthic organisms and increasing alpha diversity (Pacciardi et al. 2011E,N). The alkaloid of C. cylindracea, caulerpenyne, represents various biological activities such as antioxidant (Cavas and Yurdakoc 2005), antifungal and antibacterial properties (Vairappan 2004; Vieira Macedo et al. 2012), and antinociceptive and anti-inflammatory activities (Souza et al. 2009); several patents on Caulerpa preparations, extracts and purified metabolites have been approved (Cavas and Pohnert 2010).

Caulerpa taxifolia [Chlorophyta]: Its rapid spread, high growth rate, and its ability to form dense meadows on any type of sublittoral substrate leads to a drastic impoverishment of the algal components of the native communities and to the formation of homogenized microhabitats (Santini-Bellan et al. 1996N). It reduces species richness of native hard substrate algae by 25–55%, and, under certain conditions, outcompetes the native seagrasses Cymodocea nodosa and Posidonia oceanica or impacts their performance (de Villèle and Verlaque 1995N; Ceccherelli and Cinelli 1997E; Dumay et al. 2002N; Piazzi and Cinelli 2003N; DAISIE 2013). No negative effect of C. taxifolia on C. nodosa was found by Ceccherelli and Sechi (2002)E, who suggested that the two species are likely to co-exist in the long term. The alga’s dense clumps of rhizomes and stolons can modify quality and intensity of physical, chemical and hydrodynamic factors (Eyre et al. 2011), number and quality of shelters, quantity and quality of food to fish feeding on benthic invertebrates, and exchange of individuals between communities and populations (Santini-Bellan et al. 1996). C. taxifolia produces caulerpenyne, a toxic secondary metabolite protecting this macroalga against epiphytes and herbivores, and thus it does not provide the same trophic support as seagrasses and native seaweeds. Caulerpenyne has been associated with anti-cancer, anti-proliferative, anti-microbial, anti-herpetic and anti-viral properties; several patents on Caulerpa preparations, extracts and purified metabolites have been approved (Cavas and Pohnert 2010). Ecosystem services provided by the biotopes degraded due to C. taxifolia invasion (communities of sublittoral algae, coralligenous communities, seagrass meadows), such as food, biotic materials, coastal protection, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are negatively impacted. However, it seems that the impact of the species on native seagrass beds has been overstated; dense and healthy seagrass beds are probably not affected by C. taxifolia (Jaubert et al. 1999N, 2003; see also Glasby 2013 for related experiments in Australia, and references therein). Hence, the formation of C. taxifolia beds on already degraded seagrass beds, stressed environments, or over bare soft substrata might have an overall positive effect on some ecosystem services such as protection of sediments from erosion (Hendriks et al. 2010E).

Codium fragile subsp. fragile [Chlorophyta]: This Japanese green alga, previously referred to as Codium fragile subsp. tomentosoides, is among the most invasive species worldwide and has spread dramatically during the last century in many areas (but see also below about later declines). It alters benthic communities and habitats, its dense fronds hinder the movement of large invertebrates and fish along the bottom, and increases sedimentation (DAISIE 2013). It is a robust, large, sponge-like alga, and can crowd out and shade other plants and algae having detrimental effects on subtidal communities. Ecosystem services provided by many sublittoral biotopes, especially communities of sublittoral algae on rocky bottoms and biogenic reefs, i.e. food, biotic materials, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are potentially negatively impacted. The first record in Europe is from Ireland in 1845 (Provan et al. 2008), and it has since spread to most European Atlantic and Mediterranean coasts (Wallentinus 2007a) as well as to other continents. In several areas it has become a real nuisance, not the least due to its capacity to reproduce asexually, either by mats of vacherioid fine-branched filaments or parthenogenetic gametes (e.g. Chapman 1999; Bulleri et al. 2007 and references therein; Scheibling and Melady 2008). The species has a serious negative impact on aquaculture and shellfish fisheries: it fouls shellfish beds, smothering mussels and scallops, clogging scallop dredges, and interfering with harvesting (e.g. Neill et al. 2006; Streftaris and Zenetos 2006; Schaffelke and Hewitt 2007 and references therein). In North America, it is often called “oyster thief”, because it overgrows and smothers oyster beds (Trowbridge CD 1999 in Naylor et al. 2001), while in Europe this name refers to the introduced brown alga Colpomenia peregrina. In eastern Asia the species is cultivated for food production (e.g. Hwang et al. 2008) and is also included in cookery


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books in the western world. Ortiz et al. (2009) showed that it has a high nutritional and functional value in diets for humans and animals. The high uptake rates of especially nitrogen make it a suitable candidate, especially during the warm summer months, as a biofilter in Integrated Multi-Trophic Aquaculture (Kang et al. 2008); however, there is no such application in Europe. Since it produces DMSP (see more below) it can also have a positive impact on climate regulation (but negative on air quality). In comparison to kelp species it has relatively fast uptake rates of nutrients, implying a negative impact on competitors. If occurring in high abundances and detached, it may drift up on beaches and decompose, having a negative impact on aesthetic values and recreation and tourism. Basic physiological studies of enzymatic activities in the chloroplasts have been performed (Yu and Pedersén 1989). In the Mediterranean and off the coast of Maine (USA), observations indicate that the abundance of C. fragile subsp. fragile peaked about a decade after its first discovery, later to decline with less impact (Schaffelke and Hewitt 2007 and references therein). A similar decrease was measured lately on the Canadian east coast between 2000 and 2007, when it had declined as a dominant canopy alga from occurring in 54% of surveyed sites to 15%, while the earlier reduced native kelp had increased during the same period (Watanabe et al. 2010). Furthermore, Trowbridge and Farnham (2009) conducted field surveys in rocky pools and emergent rocky substrate for species of the genus Codium (introduced and native) during 2002–2005. They concluded that at most sites surveyed in the UK and in the Channel Islands, native Codium spp. were equally or more abundant and widely distributed than the invasive C. fragile subsp. fragile, and that no displacement or elimination of native congeners has occurred. A reduction in abundance between the early 1960s and 1997 was also reported from the Swedish west coast (Johansson et al. 1998). Positive effects of C. fragile subsp. fragile on the recruitment and survival of mussels have been reported from the northern Adriatic Sea, and its presence is likely to speed up the process of recovery of mussel beds after their integrity has been disrupted by disturbances from heat stress or human harvesting (Bulleri et al. 2006E,N and references therein). Experiments showed that bay scallops could survive and grow among C. fragile subsp. fragile in declining seagrass beds outside New York (Carroll and Peterson 2013 and references therein). There have been few grazing studies along the NE Atlantic shores, which probably is due to the fact that it is no longer such a dominant alga; also, in UK, since 1981, it is forbidden to perform field experiments with this alga (Trowbridge and Farnham 2009). From the Canadian east coast, C. fragile subsp. fragile was shown (Lyons et al. 2007) to contain relatively high concentrations of dimethylsulfoniopropionate (DMSP) and the products of its cleavage, dimethylsulphide (DMS), and acrylic acid (AA). The two latter act as chemical defence against the sea urchin Strongylocentrotus droebachiensis, and in feeding experiments the species was not eaten when other algae were available. In the same area, the periwinkle Littorina littorea also avoided plants larger than 3 cm (Scheibling et al. 2008), but by grazing germlings could limit their abundance in high intertidal pools. It was shown by Krumhansl and Scheibling (2012), by measuring nutritional quality and isotopic composition in detritus from C. fragile subsp. fragile, that the transfer of nutrients within, and between, trophic levels was altered in comparison to detritus from the native kelp Saccharina latissima (no such studies performed in Europe). Surveys of epiphytic algae in Rhode Islands (Jones and Thornber 2010) showed that C. fragile subsp. fragile was important as an ecosystem engineer with the largest number of epiphytes, especially during the cold season. This species also provides food for many native herbivorous species (ME Çinar, pers.obs.).

Gracilaria vermiculophylla [Rhodophyta]: This perennial red alga arrived in Europe most likely from the East China Sea/Sea of Japan (Kim et al. 2010). It is often loose-lying on soft substrate and can tolerate low salinity conditions and persist even under low light conditions (Rueness 2005; Nyberg 2007; Weinberger et al. 2008; Nejrup and Pedersen 2012). Based on the specific traits of this species (e.g. a remarkable ability to recruit from spores as well as from fragments and a high tolerance to environmental stress), Nyberg (2007) placed G. vermiculophylla among the 4 most potent invaders out of 114 nonindigenous macroalgal species in Europe, using the same criteria as in Nyberg and Wallentinus (2005). It is often found entangled in fishing gears and can grow on shells (Thomsen et al. 2007), thus a negative impact on fishery. On the other hand, there is a positive impact in that G. vermiculophylla attracts many mobile invertebrates eaten by fish (Nyberg et al. 2009), and on barren muddy bottoms acts as a shelter and nursery area (Weinberger et al. 2008). In the Chesapeake Bay area, E USA, it provides the blue crab with a new nursery area and thus can compensate severe declines in seagrasses (Johnston and Lipcius 2012). It is an agarophyte, now being farmed as a biofilter in an Integrated Multi-Trophic Aquaculture facility in Portugal (Abreu et al. 2011aE and references therein), and is used for agar


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production (Sousa et al. 2012 and references therein). It is also used as a biofilter, when farming mussels during the warm season in eastern Russia (Skriptsova and Miroshnikova 2011E). The large plants, when lying loose on soft substrate, trap much sediment, and also prevent resuspension, making the water more clear (Sfriso et al. 2012). The large-sized and persistent thalli also function as a carbon sink, even though competition with seagrasses may be a negative impact for the total outcome (cf. Figueiredo da Silva et al. 2009) of this process, as well as for coastal protection. The drifting plants are often driven up on the beaches, a negative impact on aesthetic values and on recreation. Basic studies of ecophysiological processes of G. vermiculophylla have been carried out, e.g. in relation to light and photosynthesis (Phooprong et al. 2008; Roleda et al. 2012) and nitrogen uptake (Abreu et al. 2011b). Analyses of samples of G. vermiculophylla from the Kiel Fjord, N Germany, showed that the epibacterial communities were quite different from those on the co-occurring native macroalgae Fucus vesiculosus and Ulva intestinalis (Lachnit et al. 2011), affecting ecosystem functions. They also reported that the epibacterial communities of G. vermiculophylla harboured several bacterial strains within the Rhizobiales, Actinobacter and Roseobacter clades, which also appeared in the microbial consortia of the red alga Delisea pulchra (Bonnemaisoniales); bacterial groups known for their antilarval or antibacterial activity. In Portugal, beds of the red-listed seagrass Zostera noltii can be converted to monospecific stands of G. vermiculophylla, which resulted in a higher productivity of the area, but lower photosynthetic efficiency (Cacabelos et al. 2012N and references therein). Also, in lagoons in northern Italy, it can replace all other macroalgae, and thus has a negative impact on their life cycles (Sfriso et al. 2012). Negative impacts on the biomass, metabolism and survival of Zostera marinahave been found (Martínez-Lüscher and Holmer 2010E; Thomsen et al. 2013E), especially at high temperatures (Martínez-Lüscher and Holmer 2010E); no such impact was found by Höffle et al. (2011)E. Ecosystem services provided by many sublittoral biotopes, especially communities of sublittoral algae on rocky bottoms and biogenic reefs, i.e. food, biotic materials, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are potentially negatively impacted. Only few grazers, even though they were able to eat G. vermiculophylla, preferred it as food, thus a negative impact, when it comes to how the system works, if it outcompetes other macroalgae (Weinberger et al. 2008; Nejrup et al. 2012). Gulbransen and McGlathery (2013) reported, by using the isotope15N, that in coastal US ecosystems, sediment on marshes and mudflats, marsh cordgrass (Spartina alterniflora), and mudflat invertebrates all incorporated nitrogen from G. vermiculophylla, indicating that this perennial alga is important (no similar studies have been carried out in Europe) in the transfer of nutrients within, and between, the trophic levels of these ecosystems (cf. also Thomsen et al. 2009). However, by building up thick mats, it drastically reduces light and nutrient availability for other macroalgae. On the other hand, G. vermiculophylla is an ecosystem engineer, creating new habitats for a suite of associated epiphytes and invertebrates, and in previously non-vegetated soft-sediment estuaries, the invasion of G. vermiculophylla may lead to an increase in abundances of native invertebrates and epiphytic algae, with likely cascading effects on higher trophic levels (Nyberg et al. 2009; Thomsen et al. 2010E, 2013E; Thomsen 2010E).

Grateloupia turuturu [Rhodophyta]: This foliose and irregular, very large, Japanese red alga most probably arrived via imported Japanese oysters (Cecere et al. 2011), on the east Atlantic and Mediterranean coasts, where it previously was misidentified as Grateloupia doryphora (Gavio and Fredericq 2002). It has a negative impact by fouling shellfish and aquaculture facilities in coastal lagoons, as well as by growing on fishing nets (Streftaris and Zenetos 2006; Cecere et al. 2011). The species is edible, especially rich in dietary fibres, but also rich in proteins and with a high percentage of polyunsaturated fatty acids; algae collected throughout the seasons in N France were analysed to see when it was best to collect samples for food use (Denis et al. 2010; Kendel et al. 2013). The species is also studied for the best way to extract phycoerythrin that can be used for both cell biological and immunological purposes, but also as a dye in the cosmetic industry and in natural food (Denis et al. 2009a,b). Chinese scientists have studied the life cycle in detail to be able to scale up the artificial seedling cultures of G. turuturu, since most of the material used today comes from wild plants (Wang et al. 2012). The large size implies it binds a lot of carbon, which, however, is mineralized when the thalli deterioriate during the warm season. Extracts from the thalli were found to contain anti-microfouling substances, against bacteria and fungi (future potential benefit for antifouling) (Plouguerné et al. 2008). Hellio et al. (2004) showed that two substances in G. turuturu inhibited the settlement of cyprid larvae of Balanus amphitrite. Cognitive benefits have been assessed through basic physiological research, in


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which Goulard et al. (2004) showed that the species has a robust circadian rhythm for photosynthesis. G. turuturu in N France often have few epiphytic organisms (Hellio et al. 2004). On the east coast of USA, it reduces abundance and species richness of invertebrates, as well as decreases biomass of macrophytic epiphytes in comparison to the native Chondrus crispus (Jones and Thornber 2010; Janiak and Whitlatch 2012). Both the alien G. turuturu and the native C. crispus have a similar kind of life cycle with a tolerant perennial crust, which gives them a competitive advantage over other species, by monopolizing substrate availability (Cecere et al. 2011; Janiak and Whitlatch 2012 and references therein). The very large thalli occupy space and reduce light conditions for understorey algal species, and have few epiphytes on them (Jones and Thornber 2010).

Lophocladia lallemandii [Rhodophyta]: This Indo-Pacific filamentous red alga, described as Dasya lallemandii from the Red Sea, was first recorded from Europe in the Saronikos Gulf, Greece, in 1908 (Athanasiadis 1987), and considered as a Lessepsian immigrant (Ballesteros et al. 2007 and references therein). It has since spread to the W and NW Mediterranean basin (Patzner 1998; Ballesteros et al. 2007N; Bedini et al. 2011). This red alga has a high invasive potential and is able to cover a variety of substrata (bare bedrocks, rocky reefs with macrophytic communities, Posidonia oceanica meadows, and coralligenous communities), becoming dominant, displacing key species, and giving the benthic seascape a homogeneous appearance (Boudouresque and Verlaque 2002; Ballesteros et al. 2007N; Cebrian and Ballesteros 2010). It can form dense, 5–6 cm high, extensive mats, which the seagrass leaves cannot penetrate, with a cover of up to 100% and occurring down to 65 m depth (Patzner 1998; Ballesteros 2006). L. lallemandii invades P. oceanica meadows causing a decrease in size and weight of the seagrass shoots, leaf chlorosis, leaf necrosis, and shoot death (Ballesteros et al. 2007N; Sureda et al. 2008N). The potential of invasion by L. lallemandii is very high inside the seagrass meadows. By trapping sediment, L. lallemandii reduces water exchange and creates anoxia within the mats (Ballesteros et al. 2007N). The modification of the microhabitat characteristics of the seagrass beds induced by the presence of L. lallemandii also affects the faunal communities associated to the seagrasses (Deudero et al. 2010N). Algae within this genus have been shown to excrete lophocladines, alkaloids with cytotoxic effects; Sureda et al. (2008)N measured the increased antioxidant response of P. oceanica, which were epiphytized by L. lallemandii in comparison to control plants, which were not. Those cytotoxic compounds were also proposed by Tomas et al. (2011a)E to be the reason why herbivorous fish avoid L. lallemandii, resulting in a negative impact on the higher levels of the foodweb. One explanation might be that no native algae produce those compounds and herbivores have not had the opportunity to evolve in order to cope with them (Cebrian et al. 2011). Furthermore, Tomas et al. (2011b)E emphasized that the commercially important sea urchin Paracentrotus lividus is negatively affected by the decline of the Posidonia-meadows caused by the dense mats of L. lallemandii, which it does not actively eat, but might reduce their abundance by eating the native macroalgae on which they are attached (Cebrian et al. 2011). By using a species of the genus from the Fiji Islands, basic research has been carried out to elucidate the chemical structure of these lophocladines and to study how they affect the cells of the impacted organisms (Gross et al. 2006). The dense mats become efficient sediment traps, a positive impact on coastal protection, although the Posidonia beds themselves are quantitatively more efficient in that process (Ballesteros et al. 2007); thus, a negative effect on this process. Dense mats of these thin filamentous plants means that they have a high surface to volume ratio, resulting in high uptake rates of nutrients and carbon dioxide (Littler 1980; Littler and Littler 1980; Wallentinus 1984), meaning a negative impact on the availability of nutrients for coarser algae. The large-scale modification of submarine landscapes induced by L. lallemandii with the overgrowth of benthic assemblages may reduce the attractiveness of the biota for underwater tourism. The thick mats, which trap sediment, have a negative impact on the aesthetic value of the ecosystem as well as on recreation. The same negative impact applies to the life cycle maintenance of other species, partly due to the reduction in diversity, but also to the trapping of sediments. This prevents other species, relying on settlement of spores, to find a solid substrate, while L. lallemandii itself can rely on vegetative fragments, which can grow out to new plants (Sureda et al. 2008). It commonly overgrows other macroalgae (Ballesteros et al. 2007N; Sciberras and Schembri 2007; Box et al. 2008N; Bedini et al. 2011) including coralline algae (Ballesteros 2006) with a negative impact on many species. It has caused a reduction in the density of the erect epiphytic bryozoan Reteporella grimaldii (Deudero et al. 2010N) and the disappearance of many benthic invertebrates, fish and macroalgae (Patzner 1998). Since L. lallemandii is growing so densely in the meadows of P. oceanica, it has a strong negative effect also on the protected native, very large bivalve,


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Pinna nobilis (Box et al. 2009N; Cabanellas-Reboredo et al. 2010N). The latter authors also showed, using stable isotopes, that there was a shift in the trophic level of the community composed by this bivalve and its associated crustaceans. Its filamentous structure means L. lallemandii provides much surface area for small epiphytic organisms. Ecosystem services provided by the biotopes degraded due to L. lallemandii invasion (seagrass meadows, communities of sublittoral algae, coralligenous communities), i.e. food, biotic materials, coastal protection, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are potentially negatively impacted.

Polysiphonia morrowii [Rhodophyta]: This filamentous red alga was originally described from S Hokkaido, Japan, and its morphology is quite similar to species native in Europe. It was first recorded in the Mediterranean Sea in southern France in 1997 (Verlaque 2001). Two years later, it was found south of Venice (Curiel et al. 2002) and there reached up to 50 cm, probably having been brought by imported shellfish or shipping. It was later identified from the coast of Brittany, France, using the barcoding technique (Geoffroy et al. 2012), showing three different haplotypes (suggesting several introductions) in that area, where it grew in extensive patches. There are hardly any documented impacts of P. morrowii on ecosystem services and biodiversity in Europe, but some types of impacts can be derived from its morphology. Since the rhizoids can attach to many different substrates, besides rocks and macroalgae, also ropes, wood, mussels and crabs (Kim et al. 2004), it probably can have a negative impact by fouling in aquaculture (Streftaris and Zenetos 2006 and references therein) or with a size of up to several dm entangle in fishing gear. Its rapid growth, due to a comparatively high ratio of surface to volume, gives them a greater potential for rapid uptake of nutrients (Littler 1980; Littler and Littler 1980; Wallentinus 1984), a negative impact for ocean nourishment. Methanol extracts and two isolated bromophenols of P. morrowii have been shown by Kim et al. (2011) to have a potential as therapeutic agents against fish viral diseases such as infectious hematopoietic necrosis virus (IHNV), infectious pancreatic necrosis virus (IPNV) and flounder spleen cells (FSP). Medical research has shown that methanol extracts have photo-protective effects against ultraviolet B radiation-induced keratinocyte damage (Piao et al. 2012b). The relatively large size indicates that it may reduce light for other algae, but filamentous algae also provide much surface for small organisms.

Sargassum muticum [Ochrophyta]: This brown alga from the NW Pacific may reach a length of several metres. There are many reports from several areas on the negative impact of this species, especially on primary producers, within the first decade after its introduction in Europe in 1971, after which it spread rapidly along the Atlantic and later the Mediterranean coasts (see e.g. Wallentinus 1999, 2002; Strong et al. 2006N; Harries et al. 2007N; Josefsson and Jansson 2013 and references therein). Based on three different molecular markers, Cheang et al. (2010) showed that there was no genetic variability in material from the introduced areas, confirming that there is no cryptic diversity there, nor in most part of Japan. Problems associated with this Japanese seaweed include replacement of native species, increase of filamentous epiphytic algae, changes in composition of flora and fauna, increased sedimentation, interference with coastal fisheries, large accumulations of drift algae, blocking of narrow sounds and harbours, and interference with recreational activities (Critchley et al. 1990; Josefsson and Jansson 2013).

S. muticum is a strong competitor with native flora for space and light through its fast growth, high fertility with germlings attached before being released, high biomass, and high densities, which can prevent settlement and development of other algae (Critchley et al. 1986; Staehr et al. 2000; Steen and Rueness 2004; Schaffelke and Hewitt 2007). It has been reported to dominate the macroalgal assemblages of many sites and has caused the decline of many seaweeds and seagrasses, such as Antithamnion plumula, Elachista fucicola, Laminaria saccharina, Laminaria digitata, Fucus vesiculosus, Gelidium spinosum, Codium fragile, Cystoseira barbata, Halidrys siliquosa, Macrocystis pyrifera, Polysiphonia nigrescens, Rhodomela larix and the eelgrass Zostera marina (Viejo 1997E; Cosson 1999; Staehr et al. 2000; Wallentinus2002; Occhipinti Ambrogi 2002; Sánchez et al. 2005N; Lang and Buschbaum 2010E). Such impacts seem to be more intense on subtidal than on intertidal assemblages (Sánchez et al. 2005N; Buschbaum et al. 2006N; Olabarria et al. 2009E). Ecosystem services provided by the affected biotopes, i.e. food, biotic materials, coastal protection, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are potentially negatively impacted.

S. muticum is a habitat engineer, as it causes substantial changes to the invaded habitats. The morphology of the species, with its lateral branches lifted in an upright position due to the many small


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air-bladders, contributes to establishing a three-dimensional habitat for many epiphytic algae and smaller animals. Dense stands of S. muticum may increase the abundance or standing stock of epibiota or enhance epibiota diversity, provide protection for animals, and attract settlement of invertebrates, which may attract fish and other predators (Wallentinus 1999; Bjærke and Fredriksen 2003; Wernberg et al. 2004N; Sánchez et al. 2005N; Buschbaum et al. 2006N; Nohrén and Odelgård 2010E). In the Wadden Sea, more than 80 species of algae and invertebrates were found associated with this seaweed, including species that otherwise are rare or absent in the Wadden Sea. However, the results differ depending on the recipient communities. No significant impacts of S. muticum on intertidal macroalgal assemblages in northern Spain were found by Sánchez and Fernández (2005)E. On the other hand, S. muticum reduces Photosynthetically Active Radiation (PAR) with repercussions on the underlying layers, leading to a decrease in species number and surface cover (Curiel et al. 1998). It may also have physical effects on its local environment, such as effects on sedimentation and light penetration (Rueness 1985), water movement, and oxygen levels. Anoxia and development of hydrogen sulphide caused by stagnation of water movement in dense stands of S. muticum have been observed, even where natural water movement is high. Modification of soft substrate ecosystems can also be achieved by “stone-walking”, i.e. the plants drift away with small stones or molluscs and then settle further away (Strong et al. 2006N and references therein). Because S. muticum also undergoes a faster and more complete decomposition than that of the native flora it has replaced, it can increase the turnover and regeneration of nutrients and thus alter the nutrient cycle (Pedersen et al. 2005). S. muticum is able to colonise soft sediments, mainly attaching to shells and pebbles. The presence of the native eelgrass Zostera marina may aid its attachment by trapping drifting fragments and allowing S. muticum to settle directly in the mud in an otherwise unfavourable environment (Tweedley et al. 2008N).

Drifting individuals or mats of S. muticum are frequently reported to cause problems for coastal and recreational fishing by becoming entangled in fishing lines and nets, as well as in boat propellers. The very long floating canopies collect both sediment and floating debris, lowering the aesthetic values (e.g. Sfriso and Facca 2013) of calm bays and canals and is often a nuisance for recreation by turning sandy bays into Sargassum beds, which often trap sediment, and senescent canopies drift up on the beaches. Rotting mats washed ashore may cause a strong odour, which is considered a nuisance and an eyesore. S. muticum can also cause clogging of intake pipes to waterworks and industrial plants using seawater for cooling. It also causes many problems in aquaculture as it fouls aquaculture structures such as cages and ropes (e.g. Verlaque 2001; Boudouresque and Verlaque 2002; Kraan 2008; Lang and Buschbaum 2010 and references therein) and sometimes drifts away with live molluscs, an important vector for spreading (Mineur et al. 2007 and references therein). Dense growth of S. muticum in or near aquaculture facilities may reduce light and impede water circulation, and thus limit oyster growth. Kraan (2008) also expressed concern that, if it is growing together with the native, commercially important and industrially harvested brown alga Ascophyllum nodosum, it may lower the value of the processed end-product. By changing the intertidal algal communities in Ireland, it may have negative implications for seaweed harvesters in rural areas and their traditional way of living. It has a potentially positive value for the extraction of polyphenols (future potential benefit), and had the highest amount of those substances among six brown algae analyzed (Jard et al. 2013), while the amount of alginate was the lowest. It is also possible to combine hydrothermal fractionation with the extraction of other valuable substances with antioxidant properties (González-Lopéz et al. 2012 and references therein). When growing in sheltered shallow areas, the long canopies often interlock, preventing water exchange and accumulating sediment, a negative impact on water quality and often the stagnant water can get very warm (Strong et al. 2006N). Sfriso and Facca (2013) proposed that in Venice, like the introduced brown alga Undaria pinnatifida (for which data were available), it accumulated PAHs and probably also petroleum compounds spread on the water surface, a positive impact on water purification. The large thalli, with a fast growth rate (Pedersen et al.al., 2005), store many nutrients, a negative impact on ocean nourishment, although the relative fast decomposition of detached annual lateral shoots makes them less efficient in that context than other perennial, large brown algae. Extracts of S. muticum have been found to contain anti-microfouling substances, against bacteria and fungi (Plouguerné et al. 2008). When dichloromethane extracts were painted and tested in situ for two months, they were more efficient against barnacles, mussels and algal spores than paints with only copper and no cytotoxicity was observed (Bazes et al. 2009); thus, a potential for being used in biological regulation. Several aspects of basic research have been carried out with S. muticum, i.e. having cognitive benefits. In ecology, Yun et al. (2012) studied how water-borne


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cues from isopods induced changes in this species and in native brown algae and found that the effects of this alien species mediated grazer-deterrent responses in native seaweeds, but not vice versa. There are many studies on how chemically treated S. muticum biomass can be used to remove heavy metals and how these mechanisms function (Carro et al. 2013 and references therein).

Stypopodium schimperi [Ochrophyta]: This membranous, flat brown alga is considered a Lessepsian immigrant, originally described from the Red Sea coast of Egypt. In the Mediterranean Sea, it occurs from the surface down to at least 25 m depth and has lately had a rapid expansion in the Aegean Sea with very high abundances, and on Rhodes Island in 2009 occasionally monopolizing the substrate (Tsiamis et al. 2010 and references therein). Invasive behaviour has earlier been seen along the Levantine coasts (Boudouresque and Verlaque 2002). Its thin thalli with a large surface area usually have rapid uptake rates for nutrients, with a negative impact on competing algae. It functions as an ecosystem engineer, it can foul submarine infrastructure, and seasonally huge quantities are cast ashore on the Syrian beaches (Marc Verlaque, pers. comm. as quoted by Streftaris and Zenetos 2006). The latter means a negative impact on aesthetic values and tourism. If the substrate is monopolized, this implies a negative impact on other sessile organisms. Habitat modification could be either positive, by creating substrate for small epiphytes, or negative by occupying space. However, targeted studies to assess its impact on Mediterranean ecological components and ecosystem services more in detail are lacking. The chemical structure of a new phenolic lipid, schimperiol, has been identified (Sampli et al. 2000) and a photosystem Q(B) protein from S. schimperi has been sequenced for further studies on how biological processes are regulated (http://www.uniprot.org/uniprot/A7J2E9).

Ulva australis (ex U. pertusa) [Chlorophyta]: This green alga is a highly competitive seaweed with high rates of nutrient assimilation and propagule production. Couceiro et al. (2011) showed that the populations in Malaga, S Spain, have arrived through an independent event compared to those in NW Spain. U. australis is not generally dominant within the Ulvales communities in European waters, which, in the Mediterranean Sea, consist mainly of U. rigida and U. rotundata (Baamonde López et al. 2007). In its native range, U. australis has several uses: as a biofilter, as a natural food in gastropod culture, in the production of processed single-cell detritus food for gastropod culture, in human nutrition as a dietary supplement with hypocholesterolaemic properties (Baamonde López et al. 2007 and references therein; CABI 2013 and references therein). In European waters, no impact on ecosystem services and biodiversity has been reported.

Undaria pinnatifida [Ochrophyta]: This brown alga, described from Japan and native in the NW Pacific, has since the early 1970s spread to all continents except Africa and Antarctica. It has a remarkable ability to rapidly colonize a wide range of substrata, including artificial ones, oyster and mussel beds, and disturbed areas. The sporophytes can grow very fast, reaching lengths of up to 2–3 metres, have high physiological tolerance to adverse conditions, and the nearly invisible gametophytes are able to survive out of water for more than a month and act as a seed bank (Fletcher and Farrell 1999; Wallentinus 2007b). U. pinnatifida has a negative impact on shellfish aquaculture, as it is known to foul live molluscs, and aquaculture structures (Wallentinus 2007b and references therein). On the other hand, U. pinnatifida has an economic value as a source of food (“wakame”), which has been the motivation for intentional introductions to some areas for farming (northern France, northern Spain) (Wallentinus 2007b and references therein; Peteiro and Freire 2012 and references therein). By using introduced U. pinnatifida from New Zealand, Mak et al. (2013) found that fucoidan could be a resource for producing natural antioxidants (future potential benefit for biotic materials), and char produced by fast pyrolysis of this alga can be used as an adsorbent for copper from aqueous solutions (Cho et al. 2013). Growing plants function as a biofilter when farming mussels during the cold season (Skriptsova and Miroshnikova 2011E). U. pinnatifida accumulates both organic pollutants and inorganic metals from the polluted water in Venice (Sfriso and Facca 2013 and references therein). Since it grows on hard substrate, including artificial substrate, and has a flat morphology, it does not contribute to the protection of the coasts. The flat and comparatively thin thalli have quite rapid nutrient uptake rates, which means that competing with coarser species can be a disadvantage. Fletcher and Farell (1999) pointed out that, in areas with high sediment load and lower salinities, where less native vegetation occurs, U. pinnatifida may even be beneficial by providing a nursery ground for small fish and shelter for macrofauna, and in Argentina it was found to increase diversity and species richness of macrofauna, both mobile and sessile species, by creating a more complex habitat (Irigoyen et al. 2011b). From the surface of the thallus of U. pinnatifida, Hong and Cho (2013) isolated the epibiontic bacterium Streptomyces violaceoruber, from


15

which they isolated two bioactive furanone derivatives displaying significant anti-fouling activity (future potential benefit for antifouling). Another species of the same bacterial genus, isolated from the hapters of U. pinnatifida, produced two bioactive benzaldehydes, which were shown to kill bacterial fish pathogens (potential future benefit for biological regulation) (Cho and Kim 2012). In NW France, large amounts of detached plants were seen cast ashore on beaches in the early 1990s which reduces the aesthetic quality of the shores (Hay and Villouta 1993: 475) and has a negative impact on recreation. In Argentina, the species has been reported to negatively affect a number of recreational (diving, angling, spear fishing, fouling on boats etc.) and commercial activities (Irigoyen et al. 2011a and references therein). There are many papers on cognitive benefits, especially for medicine (e.g. Yoo et al. 2012), products for functional food (e.g. Piovan et al. 2013 using algae from Venice) and basic physiological science (e.g. Qiao et al. 2013). In Venice, Italy, Curiel et al. (2001) reported, seven years after it was first recorded in 1992, that colonization was in progress and that a significant decrease in the surface covered by other algae had occurred, suggesting competition. However, Sfriso and Facca (2013) performed monthly samplings in the lagoon ten years later and found that the total stand stock of this species was about 0,2 ktonnes FWT y-1 in the entire lagoon, but that, together with the introduced Sargassum muticum (with 25 times higher biomass), it only constituted 5-7% of the total macroalgal biomass there. In most of Europe, where it has been introduced, the trend has been for a negligible or small negative impact on indigenous macroalgae. The invasiveness of U. pinnatifida is due, to a large extent, to the frequency of vectors spreading it to new localities (cf. Hay 1990), since in several areas it is dependent on gaps in the communities of large canopy organisms, which this annual plant, with a very high release of zoospores and tolerant gametophytes, can occupy quickly by its opportunistic growth pattern, although the native species might return after the detachment or degeneration of the Undaria sporophytes (for references see Wallentinus 2007b; Schaffelke and Hewitt 2007; Schiel and Thompson 2012; Thompson and Schiel 2012). It has been a food item for different grazers in several areas (Wallentinus 2007b: Table 4.7.1). From Argentina, Irigoyen et al. (2011a) reported that drifting detached U. pinnatifida thalli physically obstructed access to shelters in the reef for several fish species, and the large size of the thalli have a negative impact on smaller algae by shading. However, the large thallus area means that it creates new habitats for many epiphytic organisms and nursery ground for mobile animals (see references in Wallentinus 2007b).

Womersleyella setacea [Rhodophyta]: This turf-forming red alga, described from the Hawaiian Islands as Polysiphonia setacea, originally has an Indo-Pacific-Caribbean distribution, and colonizes wide zones throughout the Mediterranean Sea, forming thick persistent carpets that completely cover deep sublittoral rocky substrata and seagrass measows (Athanasiadis 1997; Airoldi et al. 1994; Airoldi 2000E; Piazzi and Cinelli 2001, 2003N; Streftaris and Zenetos 2006; Battelli and Rindi 2008; Cebrian and Rodríguez-Prieto 2012). The persistence of these carpets over the year results in a stable habitat for many animals, which sometimes increase in both abundance and number of species (Antoniadou and Chintiroglou 2007; Cebrian and Rodríguez-Prieto 2012 and references therein). It acts as a habitat/ecosystem engineer (Wallentinus and Nyberg 2007; Berke 2010) by negatively affecting the available substrate for other epiphytic macroorganisms to settle on, and by reducing the light for other autotrophs, with substantial negative effects on native communities; it modifies benthic assemblages, reduces diversity, affects food webs, and outcompetes key species (Piazzi and Cinelli 2000, 2001, 2003N; Serio et al. 2006; Battelli and Rindi 2008; Piazzi and Balata 2009N; Cebrian and Rodríguez-Prieto 2012; Linares et al. 2012E; Piazzi et al. 2012; de Caralt and Cebrian 2013N). On the other hand, it may provide more surface area for microorganisms to grow on than the seagrass plants do, and in some areas W. setacea can be the only dominant species in its functional group increasing the structural complexity (Antoniadou and Chintiroglou 2007). Together with Acrothamnion preissii, they have an adverse impact on the protected Posidonia oceanica meadows, reducing the functional diversity of the epiphytic macroalgal community of the rhizomes and occupying about 50% of the macroalgal cover on dead P. oceanica mattes (Piazzi and Cinelli 2000N). When there was a dense turf of W. setacea, it was noted that P. oceanica had a reduced number of leaves per rhizome (Battelli and Rindi 2008). Most herbivores do not graze it (Cebrian and Rodríguez-Prieto 2012 and references therein), with a negative impact on the food web. It also impacts coralligenous communities, as it has been found to form a monospecific layer covering the coralligenous substrata (Ribera and Boudouresque 1995; Piazzi et al. 2012). It has a negative effect on the assemblages associated with maerl beds, as it substantially reduces the amount of photosynthetically active radiation reaching the photosynthetic tissues of the rhodolith-forming algae,


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and traps sediment that may create an anoxic boundary-layer around the thallus of the rhodolith-forming algae, smothering them and limiting rhodolith growth (Sciberras and Schembri 2007). It has a strong and consistently negative effect on various components of fitness of the endangered slow-growing and long-lived gorgonian Paramuricea clavata (Cebrian et al. 2012E). Reduced recruitment and increased mortality of gorgonian juveniles (Linares et al. 2012E) and a strong negative impact on the reproduction of several deep-living sponges (de Caralt and Cebrian 2013N) have been reported. It reduces the diversity of the seagrass meadows and can clog fishing nets (Verlaque 1989). The turf efficiently traps sediment (Piazzi and Cinelli 2001), a positive impact on coastal protection, although the Posidonia beds themselves are quantitatively more efficient in that process (Ballesteros et al. 2007). The trapping of sediments has a negative impact on the aesthetic value of the ecosystem as well as on recreation. The same negative impact applies to the life cycle maintenance of other species, partly due to the reduction in diversity, but also to the trapping of sediments. This prevents other species, relying on settlement of spores to find a solid substrate for settlement, while W. setacea itself totally relies on vegetative fragments, which can grow into new plants (e.g. Piazzi and Cinelli 2001; Piazzi and Balata 2009N and references therein). Airoldi (2000)E showed that benthic crusts can coexist with turf of this species, although the negative impact increases when other stressors occur simultaneously (Piazzi et al. 2012 and references therein). Dense turf of these thin filamentous plants means they have a high surface to volume ratio, resulting in high uptake rates of nutrients and carbon dioxide (Littler 1980; Littler and Littler 1980; Wallentinus 1984), resulting in a negative impact on the availability of nutrients for coarser algae. Ecosystem services provided by communities of sublittoral algae and seagrasses degraded due to W. setacea, i.e. food, biotic materials, coastal protection, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are negatively impacted. It has not been possible to find any cognitive benefits for this species.

Tracheophyta

Halophila stipulacea: This seagrass species (type locality in Yemen, southern Red Sea) entered in the Mediterranean through the Suez Canal, possibly transported by boats or fishing gear (Lipkin 1975). It has been reported to outcompete the native Mediterranean seagrasses and induce changes in the sublittoral communities (DAISIE 2013); however, we found no evidence for such an impact. On the other hand, the rapid colonization of recently disturbed habitats (e.g. due to storms) by H. stipulacea could interfere with natural seagrass succession. The fish community in the meadows of eastern Sicily was observed to be similar to meadows of other Mediterranean seagrasses, which have a stable structure over the year (Di Martino et al. 2007), and many small and juvenile littoral species were present, pointing to their importance as nursery areas. Also in the Caribbean, where it was first seen in 2001 (Ruiz and Ballantine 2004), a rich fish community was revealed in H. stipulacea beds (Willette and Ambrose 2012). Being a rooted plant it can utilize nutrients in the sediments, which are then transferred to the foodweb by grazers (e.g. Mariani and Alcoverro 1999). A preliminary screening showed that acetone extracts had an insecticidal activity, exhibiting high mortality against rice weevil, and in the future could be developed to replace chemical compounds (Gnanambal and Patterson 2006). Phytochemical analyses have revealed new natural compounds and their chemical structures (Bitam 2011). Also, growing down to a depth of almost 50 m in its native area, there are several basic photoacclimation studies using H. stipulacea (e.g. Sharon et al. 2011). In comparison with native Mediterranean seagrasses, H. stipulacea was found to have a poor epiphytic algal flora and coralline crusts were virtually missing (Rindi et al. 1999), the hypothesis being the fast turnover rate of the leaves. When growing on previously barren areas as a pioneer species (Mariani and Alcoverro 1999) it provides new architecture and shelter for the mobile fauna. H. stipulacea beds, like all seagrass ecosystems, provide a great variety of services; they provide habitat, shelter, food source and nursery grounds for a variety of marine organisms, enhance local biodiversity, oxygenate waters and sediments, contribute to cycling nutrients, act as net carbon sinks, control the transparency of the water column by favouring retention of suspended particles, and protect shorelines by their networks of rhizomes that stabilize sublittoral sediments (Salomidi et al. 2012). Hence, a higher cover of H. stipulacea in contrast to bare sediments may overall be beneficial for many ecosystem services.


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Spartina alterniflora: S. alterniflora is a rhizomatous, perennial salt marsh grass that can form extensive monoculture meadows. Many coastal engineers used it, together with other Spartina species, to control natural coastal erosion (CABI 2013). Natural coastal habitats in Europe have been altered to monoculture Spartina meadows, resulting in displacement of flora and fauna. In many areas, S. alterniflora has replaced the native S. maritima and it has been reported to menace the rare populations of Limonium humile (Campos et al. 2004). It is a habitat engineer; it has a dense rhizome/root system that binds coastal mud and its sturdy stem decreases wave action allowing silt deposition, causing elevation of the mudbank, changing water circulation patterns, reducing tidal flow, assisting in land reclamation, and replacing open mudflat habitats associated with bottom-dwelling invertebrate communities by vegetative salt march species (CABI 2013). The presence of S. alterniflora increases the oxidizing capacity of sediments and enhances total microbial mineralization in comparison to unvegetated areas (Gribsholt and Kristensen 2002E). S. alterniflora produces DMSP (e.g. Yoch 2002), with a possible positive impact on climate regulation (see the text for Phaeocystis pouchetii). On the other hand, S. alterniflora communities have been shown to produce high emissions of the greenhouse gas methane (Tong et al. 2012), a negative impact on climate. Spartina alterniflora from North America was introduced in the UK and hybridized with the local Spartina maritima, producing a sterile hybrid, S. townsendii. The hybrid subsequently underwent autogenic chromosome doubling to produce a new high-impact invasive fertile species, Spartina anglica (see below). Hence, S. alterniflora may be considered co-responsible for all the positive or negative impacts that have been reported for S. anglica as well.

Spartina townsendii var. anglica (hearafter S. anglica): It is a fertile hybrid of the European indigenous Spartina maritima and the North American Spartina alterniflora. S. anglica can accrete large volumes of tidal sediment leading to substantial increases in marsh elevation. This property made the species valuable for coastal protection, stabilization of mudflats, and reclamation schemes in the early twentieth century, and the species has been intentionally introduced to coastal and estuarine mudflats throughout the world (Gray et al. 1991; Nehring and Adsersen 2006). It was also introduced in estuarine areas for the benefit of agriculture as a wide variety of livestock (i.e. cattle, goat) will eat S. anglica (Ranwell 1967). S. anglica beds, similar to all seagrass ecosystems, provide a great variety of services; they provide habitat, shelter, food source and nursery grounds for a variety of marine organisms, enhance local biodiversity, act as net carbon sinks, and protect shorelines (Salomidi et al. 2012). S. anglica is highly invasive and can cause a loss of valuable habitat for endobenthic invertebrates and for migrating shorebirds and waterfowl and a loss of nursery areas for fish, it can displace native plants and more diverse native plant communities and alter the course of succession (Nehring and Adsersen 2006). S. anglica invades former native salt marsh grass-dominated vegetation, and is perceived as a threat to salt marsh conservation, displacing native flora, such as S. maritima, Salicornia spp. and Zostera noltii, and benthic infauna (e.g. Arenicola marina, Neries diversicolor, Corophium volutator) with consequences for some wildfowl and waders (Nehring and Adsersen 2006 and references therein). Mesocosm experiments indicate that S. anglica strongly influences the biogeochemical turnover in the sediment, possibly because of its strong ability to transport oxygen to the substratum (Gribsholt and Kristensen 2002E). It may compete with areas used for oyster farming and impede recreational activities (DAISIE 2013). Along the Kattegat coast some of the establishments occur on sandy beaches that are attractive to tourists, where formation of large swards would change the aspect of the beach and create a belt with unattractive sedimentation (Nehring and Adsersen 2006). On the other hand, S. anglica is a habitat engineer, may form the basis of many food webs and is a possible food source for many grazers such as geese, ducks and other water birds and wildlife (Nehring and Adsersen 2006). The species may have potential as a biofuel, paper and as a food product for animals (DAISIE 2013).

Polychaeta

Ficopomatus enigmaticus [Annelida]: This serpulid polychaete species is an ecosystem engineer, creating reef-like aggregates in which tubes grow vertical to the substrate in clumps and attach to each other (ten Hove 1979; Schwindt et al. 2001). This species preferably colonizes hard substrate in brackish water, but also forms reefs in tropical and subtropical areas, which have very hot summers and highly saline conditions (Pillai 2008). It might reach an annual production in dry weight of almost 21 kg.m-2.yr-1


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(Fornós et al. 1997) and a density of 150,000 ind.m-2 (Bianchi and Morri 2001). It sometimes forms isolated aggregates which are circular, reaching up to 4 m in diameter and 0.5 m in height (Obenat and Pezzani 1994). Massive F. enigmaticus aggregates in invaded lagoons can have dramatic consequences for the biophysical properties and processes as well as resident biota (Davies et al. 1989; Schwindt et al. 2001, 2004). This species feeds on suspended detritus and phytoplankton (Bruschetti et al. 2008E) and the ingestion rate was estimated at almost 45 mg mg-1dry mass of worm h-1 (Davies et al. 1989). The feeding activity of this species substantially reduces sea-water turbidity, removes suspended particulate matter, reduces phytoplankton concentrations, excretes large amounts of inorganic nutrients, and increases silt accumulation by trapping sediment between tubes (Keene 1980E; Davies et al. 1989; Bruschetti et al. 2008E; Dittmann et al. 2009). The clearance of water by this species also greatly improves light penetration, resulting in an increase in benthic productivity (Bruschetti et al. 2008E). F. enigmaticus aggregates provide substrate and shelter for many epibionts and represent excellent food for many species including fishes (Obenat and Pezzani 1994; Chuwen et al. 2007). In Emsworth millpond (England), 24 macroinvertebrate species (polychaetes, amphipods and isopods) inhabit the microhabitat between the reefs tubes (Thomas and Thorp 1994). In the Po River Delta (Italy), more than 15 macroinvertebrate species plus several algae species are associated to the reefs, the most abundant being amphipods, polychaetes, barnacles and bryozoans (Bianchi and Morri 1996). This species forms reefs (EUNIS Cod: A2.73) in the Black Sea, harbours 46 invertebrate and 3 fish species, and sustains a high settlement of associated fauna (max. density: 296,278 ind m-2, biomass: 5,885 gDW m-2) (Micu and Micu 2004). F. enigmaticus reefs can have a positive effect on the habitat use of birds because the area of reefs was preferred for feeding and resting (Bruschetti et al. 2009). F. enigmaticus colonies attach to water intake pipes, fishing boats, leisure craft, floating structures in lagoons and docks, aquaculture ponds, ports, and docks, inducing costs for cleaning and maintenance (Nelson-Smith 1967; Zaitsev and Öztürk2001; Jensen and Knudsen 2005; Muniz et al. 2005).

Hydroides dianthus [Annelida]: Hydroides dianthus is a primary fouling species that can form massive aggregates on artificial substrates such as ship hulls, quays and mariculture equipment (Relini 1993; Toonen and Pawlik 1996; Koçak et al. 1999). It builds up dense populations (e.g. 33,050 ind. m-2) in harbour environments (Çinar et al. 2008). Ficopomatus enigmaticus and H. dianthus built extensive reefs in the lagoon of Orbetello (Tuscany, Italy), forming 1-m thick layers which covered about 7% and 1.5% of the lagoon area, respectively. The maximum tube density of H. dianthus reefs was about 12,000 ind. m-2 (Bianchi and Morri 2001). As it probably feeds on bacteria and phytoplankton like H. elegans, dense populations of this species in polluted waters might contribute to water purification. The filtration rate of the congenetic species H. norvegica is about 0.011 L/h (Dales 1957). The mass calcareous structures built by H. dianthus seem to provide a suitable microenvironment for sessile (mainly hydroids) and mobile (mainly pycnogonids and syllid polychaetes) species (Haines and Maurer 1980; Çinar et al. 2008). This species is the host of the trematod Cercaria loossi which could be a parasite in fish blood (Linton 1915; Zibrowius 1978).

Hydroides elegans [Annelida]: Hydroides elegans is a primary fouling species, occurring abundantly, especially in harbour environments. Its population density can reach 110,700 ind. m-2 in the Mediterranean (Çinar et al. 2008) and 330,000 ind. m-2 in the Pacific (Jianjun and Zongguo 1993). Massive aggregates created by H. elegans provide a suitable microenvironment for other benthic species (Koçak et al. 1999; Çinar et al. 2008). As it mainly feeds on bacteria and phytoplankton (Gosselin and Qian 2000; Qui and Qian 1997), dense populations of this species in polluted waters might contribute to water purification. It competes with the alien serpulid H. dianthus for space (Bianchi 1983). It can become a nuisance when attached to hard structures such as quays, mariculture equipment and ship hulls (Jianjun and Zongguo 1993; Relini 1993). It becomes abundant on artificial substrates near thermal pollution and can clog cooling systems (Zibrowius and Thorp 1989). A mass occurrence of this species has resulted in heavy economic costs (ca. 10 billion ¥) to the oyster culture industry in Japan through fouling on their shells (Hirata and Akashige 2004; Link et al. 2009). H. elegans is an excellent model organism for experimental studies and is easily adapted for laboratory biofouling research because of its short generation time (~3 wks) and ease of propagation (Nedved and Hadfield 2009).

Hydroides ezoensis [Annelida]: It is a severe fouling organism on harbour structures and ship hulls; it can cause flotation problems for buoys and adds considerably to fouling of ships (Zibrowius 1978; Zibrowius and Thorp 1989; Jianjun and Zongguo 1993). As it probably feeds on bacteria like H. elegans, dense populations of this species in polluted waters might contribute to water purification. It also forms


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large encrustations within pipes used for the sea-water cooling systems (Thorp et al. 1987). The bulk of its massive aggregations (up to 30 cm thick) is a protected habitat for free-living and sessile invertebrates; it also provides food to fish and invertebrates (Eno et al. 1997). However, dense settlement of this species may kill native sessile species such as oysters and mussels by overgrowing (Eno et al. 1997).

Marenzelleria neglecta and Marenzelleria viridis [Annelida]: M. viridis and M. neglecta were proved to be distinct species (Sikorski and Bick 2004). As these two species were previously confused with each other or misidentified in northern Europe, they are combined here as Marenzelleria spp. These burrowing polychaetes have spread rapidly in the Baltic Sea and now dominate many soft-bottom communities, where they dwell in burrows down to 40 cm depth in the sediment (Zmudzinski 1996; Zettler et al. 1994; Daunys 1997; Olenin and Leppäkoski 1999). Marenzelleria spp. have caused significant changes to the macrobenthic communities in the last decades. Intense invasion of these species has led to almost single-species benthic communities and a multiple increase in the biomass of the macrozoobenthos, especially in areas affected by hypoxia, where the original macrozoobenthos was reduced or absent. In huge areas, Marenzelleria spp. replaced the pristine community of glacial crustacean relicts and the amphipod Monoporeia affinis (e.g. Ezhova et al. 2005; Maximov 2011). Negative impacts on populations of native species (such as the polychaete Hediste diversicolor and the amphipod Monoporeia affinis), due to competition for food and space, have been reported (Kotta et al. 2001E; Kotta and Ólafsson 2003E; Neideman et al. 2003E; Kotta et al. 2006E). After the invasion of Marenzelleria spp., the densities of the amphipod Corophium volutator, the polychaete Hediste diversicolor, and the amphipod Monoporeia affinis have dropped considerably in the Baltic Sea (Atkins et al. 1987; Essink and Kleef 1993; Zettler 1996; Kotta and Kotta 1998; Cederwall et al. 1999; Kotta et al. 2001E; Maximov 2011). Marenzelleria spp. re-circulate organic matter and nutrients deposited in deeper sediment, link benthic and pelagic subsystems, affect sediment and fluid transport processes, improve oxygen circulation and aerobic transformation of organic matter, increase benthic production, affect nutrient fluxes, and provide new microhabitats for their associated fauna (Kube et al. 1996; Leppäkoski and Olenin 2000; Kotta et al. 2001E; Çinar 2013; DAISIE 2013). The presence of Marenzelleria may lead to a marked stimulation of sulphate reduction. Marenzelleria invasion may therefore have widespread consequences for microbial pathways and nutrient dynamics in both sandy and muddy marine sediments (Hietanen et al. 2007E; Quintana et al. 2013E). Bioturbation by Marenzelleria has a potential beneficial effect on its environment by improving oxygenation of the sediments (at greater depths than any of the native species of the Baltic) and speeding decomposition of the organic matter settling on the bottom. These bioirrigation activities of dense Marenzelleria populations have a major impact on sedimentary phosphorus dynamics. This may facilitate the switch from a seasonally hypoxic system back to a normoxic system by reducing the potential for sediment-induced eutrophication in the upper water column (Norkko et al. 2012). However, in areas with sediments in which nutrients and toxic substances have accumulated over a long period, this may also have harmful effects (Olenin and Leppäkoski 1999). Marenzelleria spp. are a potential food source for demersal fish, such as plaice (Pleuronectes platessa) and flounder (Platichthys flesus) (Essink and Kleef 1993; Winkler and Debus 1997). Delefosse and Kristensen (2012)E showed its positive impact on burying seeds of Zostera marina thereby reducing seed predation and facilitating seed germination.

Establishment of any single alien species in a novel region and ecosystem creates opportunities for ecological research focused on concepts such as adaptive strategies, niche dimensions, interspecific relationships, dispersal mechanisms etc. However, these invasive species have been reported to interfere with research and monitoring e.g. in the Baltic Sea, where benthic communities have been monitored by quantitative methods since the 1910s. The results of this long-term effort, based on international sampling programmes, may be invalidated by the introduction of Marenzelleria spp. in areas where they have become dominating, utilising the space and energy resources in a different manner and rate and restructuring the food webs, previously built up by native species (Leppäkoski 2002; Villnäs and Norkko 2011).

Pileolaria berkeleyana [Annelida]: This species was common in the Canary Islands and British coasts and suffered from competition with other Pileolaria species probably originating from the Mediterranean Sea (Knight-Jones and Knight-Jones 1980; Zibrowius and Thorp 1989). It was rarely found in the Mediterranean Sea (Zibrowius 1983). There are no documented impacts on ecosystem services or biodiversity in Europe.

Spirorbis (ex Janua) marioni [Annelida]: Spirorbis marioni is known to foul artificial substrates in the harbour environments of the Mediterranean (Bianchi 1981; Knight-Jones et al. 1991; Çinar 2009). It rarely becomes abundant in some places in the Mediterranean (Zibrowius 1983; Bianchi et al. 1984; Knight-Jones et al. 1991), but extraordinary abundant in the Canary Islands (Knight-Jones and Knight-


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Jones 1980). It needs shelter or artificial substrata for its existence (Knight-Jones and Knight-Jones 1980). In ports, it might compete with other sessile species.

Crustacea

Acartia tonsa: A. tonsa competes with native copepods, especially congenerics, and may dominate zooplanktonic communities (Gubanova 2000; David et al. 2007; Aravena et al. 2009; Jensen 2010a). Gomoiu et al. (2002) reported that summer mesozooplankton in the Crimea Peninsula waters is dominated by two alien species, A.tonsa and larvae of the barnacle Amphibalanus improvisus; A. tonsa occupies mainly the ecological niche that was previously occupied by the native species Acartia latisetosa. In south-western France it is seasonally replacing A. bifilosa in low salinity waters (David et al. 2007). This was attributed to intrusion of saline water plus increased water temperature in connection with a nuclear power plant. In a Spanish estuary with relatively high salinity, A. tonsa had a negative impact on A. clausi (Aravena et al. 2009; Jensen 2010a). Due to its high abundances and grazing abilities, A. tonsa can change energy and matter flows between pelagic and benthic compartments, modify trophic structure of invaded ecosystems, and even serve as a potential biological control of algal blooms (Leppäkoski et al. 2002a). Acartia spp. (including A. tonsa) constitute significant prey for fish, but the overall effect on fish of the displacement of native species by A. tonsa has not been documented. The successful colonization of A. tonsa in the Gironde estuary had led to an increase of zooplanktonic production, which could have had a positive impact on fish and shrimp production (David et al. 2007). A. tonsa has been used to produce live feed for aquaculture organisms (Sørensen et al. 2007), e.g. turbot rearing in the Black Sea.

Amphibalanus improvisus: Filter feeding barnacles transfer organic material from the water mass to the sediment - this provides an important source of energy to the benthic environment. A. improvisus can promote the settlement success and further development of filamentous algae probably by increasing nutrient availability in benthic systems through biodeposition (Kotta et al. 2006E). A. improvisus increases the 3-dimensional surface available for associated macro and meiofauna on shallow-water hard bottoms (e.g. underwater artificial substrata) and can enhance detritus-based food chains by supplying their habitat with particulate detritus. In very low-saline waters, e.g. the Baltic Sea and the uppermost reaches of estuaries, where A. improvisus is the only barnacle, the fouling impacts may be considerable (Leppäkoski and Olenin 2000). There, A. improvisus avoids competition, may facilitate settlement of other organisms, and causes marked habitat and community structure alteration (Leppäkoski and Olenin 2000). A. improvisus is a strong competitor for space. On the other hand, in dense populations of A. improvisus, biodiversity of associated species such as chironomid larvae, ostracod and copepod crustaceans and juvenile bivalves increases compared to adjacent sites without the barnacle (Leppäkoski 1999N). Empty shells of the barnacle serve as new microhabitats for small invertebrates. It causes economic losses to industries, power plants, and shipping companies by fouling intake pipes, heat exchangers, underwater constructions and ship hulls. It also causes harm to boating and other recreational uses (sharp shells). It may have a negative impact on aquaculture by fouling mussels, oysters, and underwater installations (Leppäkoski 1999). Larvae of A. improvisus are regularly used in toxicity tests, especially in connection with antifouling substances (e.g. Dahlström et al. 2000; Sjögren et al. 2008; Toth and Lindeborg 2008).

Austrominius (ex Elminius) modestus: It can become dominant, displacing native barnacle species (Barnes and Barnes 1966; Bennell 1981; Lawson et al. 2004; Franke and Gutow 2004; Witte et al. 2010; Gomes-Filho et al. 2011; Bracewell et al. 2012; CABI 2013). Bennell (1981) reported that peaks of abundance of A. modestus occurred only when the native Semibalanus balanoides cover was low, which may have been due to competition for space. Along the shores of north-western Spain and Portugal, Barnes and Barnes (1966) described it as a severe competitor of Chthamalus in relatively quiet waters, where they found Chthamalus limited to the upper shore, due to a dense zone of A. modestus lower down. Witte et al. (2010) reported that in summer 2007 at Sylt (North Sea), abundances of A. modestus had overtaken those of S. balanoides and Balanus crenatus. In the Sylt-Rømø Bight ecosystem (North Sea), Baird et al. (2012), based on ecological network analysis (ENA), reported a decline in the number of cycles, the trophic efficiency, the ascendency ratios, and relative redundancy that was ascribed to the impact of the invasive species Crassostrea gigas and A. modestus on system organization and function.


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In the years just after its introduction into Great Britain, it was suggested that A. modestus was a potential threat to the periwinkle and oyster industries due to competition with oyster spat and as it induced additional sorting and cleaning cost for fouled shells before marketing (Bassindale 1947; Knight-Jones 1948; Crisp 1948). It has been used as a test organism in toxicity tests (Corner et al. 1968).

Callinectes sapidus: The blue crab, originally from the east coast of the Americas, has been widely recorded in all European regional seas. In the eastern Mediterranean (especially in Turkey, Greece, Syria, Lebanon, Israel and Egypt) it became very abundant and is commercially exploited and frequently sold in fish markets (Zenetos et al. 2010). The annual production of this species is about 200 tonnes/year in Turkey (Çinar et al. 2005). It has been reported to mutilate fish caught in traps and trammel nets and to tear nets (Galil et al. 2013). Competition with other crabs has been assumed for Mediterranean populations (Gennaio et al. 2006).

Caprella mutica: C. mutica can be very abundant, especially in fouling communities on artificial substrata (e.g. on boat hulls, navigation/offshore buoys, floating pontoons and aquaculture infrastructure), reaching abundances of >200,000 ind m–2 during summer and early autumn (Ashton 2007; Boos 2009E). C. mutica has the ability to displace ecologically similar native species when resource space is limited and when its density is high (Boos 2009E; Shucksmith et al. 2009E; Boos et al. 2011). Detailed knowledge on community or ecosystem level impacts of the species is still lacking (Boos et al. 2011). C. mutica can be a substantial food source for fish (Page et al. 2007). Use of the species for monitoring trace metals, especially due to punctual environmental perturbations (oil spill accidents, waste outfalls, etc.), has been suggested (Guerra-García et al. 2010).

Cercopagis pengoi: The Ponto-Caspian predatory cladoceran Cercopagis pengoi was probably brought to the Gulf of Finland in 1992 with ship traffic via the Volga–Baltic corridor and soon spread to most part of the Baltic Sea (Leppäkoski et al. 2002b). It has caused the decline of native small-sized cladocerans such as Bosmina coregoni maritima, Evadne nordmanni, and Pleopsis polyphemoides probably due to direct predation (Ojaveer et al. 2004; Kotta et al. 2006). It is also a potential competitor with young stages of planktivorous fish for herbivorous zooplankton and may affect resident zooplankton communities by selective predation (Birnbaum 2011 and references therein). Such changes may result in decreased grazing pressure on phytoplankton and enhanced algal blooms (Birnbaum 2011). On the other hand, C. pengoi is itself a very important food source for many fish, such as small herring, stickleback, smelt and bleak (Ojaveer et al. 2004; Kotta et al. 2006). Consumption of C. pengoi by smelt enhances energy transfer from the surface waters to the cold deeper water layers, which is especially important during the periods of food shortage in deep water layers (Kotta et al. 2006). Thus, the effects of C. pengoi on the pelagic ecosystem are two-sided as competing for food with other pelagic predators and reducing the efficiency of energy transfer to upper trophic levels (Kotta et al. 2006). C. pengoi attaches to fishing gear and clogs nets and trawls, causing problems and substantial economic losses for fishermen and fish farms (Leppäkoski and Olenin 2000; Birnbaum 2011).

Charybdis (Charybdis) japonica: There are no documented impacts on ecosystem services or biodiversity in European waters.

Charybdis hellerii: There are no documented impacts on ecosystem services or biodiversity in Europe. C. hellerii is of commercial value in South-East Asia (Lemaitre 1995) and has been cultivated in the Philippines; however, in the Mediterranean and Atlantic it currently does not have any market value (CABI 2013). It has not yet been shown that the successful establishment and spread of C. hellerii threatens native ecosystems, habitats or species; however, it is a territorial omnivore which often feeds on other small crabs, and thus an established population may adversely affect local crab populations (Dineen et al. 2001; CABI 2013).

Chionoecetes opilio: The species is a high value commodity for fisheries (Jensen 2010c). There are no reported negative impacts on ecosystem services and biodiversity.

Eriocheir sinensis: E. sinensis (mitten crab) is an opportunistic omnivore capable of eating a wide variety of invertebrates as well as fish eggs, algae and detritus (Dittel and Epifanio 2009). The species competes for space and food especially during mass developments, can modify habitats mainly because of its burrowing activity, decrease biodiversity, and change community structure (Olenin and Leppäkoski 1999; Dittel and Epifanio 2009; DAISIE 2013). It is considered as an ecosystem engineer due to its burrowing activity, which can result in increased erosion of dikes, river and lake embankments (Clark et al 1998; DAISIE 2013). During mass occurrences, mitten crabs may clog water intakes. E. sinensis interferes with recreational and commercial fishing and can cause considerable damage to fisheries by


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consuming netted fish and by cutting nets (Streftaris and Zenetos 2006). The entanglement of E. sinensis in fish and shrimp nets increases handling time and damages the target species within the net (Dittel and Epifanio 2009 and references therein; DAISIE 2013). Mitten crabs have been used as live fish bait, for fish meal production, as agricultural fertilizer, and for cosmetic products (Dittel and Epifanio 2009; DAISIE 2013). In Asia it is considered a delicacy, supporting an annual billion-dollar fishery (Sui et al. 2009), and in certain European regions adult crabs, caught as by-catch in inland fisheries, are sold to Asian restaurants (DAISIE 2013).

Gammarus tigrinus: G. tigrinus can become dominant in intertidal and subtidal benthic habitats and may outcompete many native gammarids, such as G. duebeni, G. salinus and G. zaddachi, altering species composition and interspecific interactions (Grabowski et al. 2006, 2007; Packalén et al. 2008; Orav-Kotta et al. 2009E; Sareyka et al. 2011E). In the northern Baltic Sea, the species has a strong potential to modify benthic community structure and functioning in the whole sublittoral zone. Here G. tigrinus appears to be competitively superior over all native gammarids (except G. duebeni). Its further expansion is likely leading to further decline of the native gammarid populations in, e.g., the Gulf of Finland (Kotta et al. 2013). In oligohaline conditions, G. tigrinus is known to eliminate native gammarids via intraguild predation (Dick and Platvoet 1996E). There is also evidence of a functional impact of G. tigrinus in impacted aquatic ecosystems, as the replacement of native species leads to a substantial decrease in the rate of leaf litter recycling (Piscart et al. 2011E). G. tigrinus has been reported to have damaging effects on fishing gear and trapped fish (Pinkster et al. 1977). Recent predictions based on ecological niche modelling point out that the species has a vast potential invasive range and may become cosmopolitan through shipping (Ba et al. 2010). The species is euryhaline and also colonizes brackish and fresh waters and effectively switches between habitats with different salinity conditions (Kelly et al. 2006; Normant et al. 2007). Thus, once established in an estuary it may also pose risks for inland waters.

Hemigrapsus sanguineus: The Asian shore crab H. sanguineus is an omnivorous predator capable of opening small mussels and probably young oysters. During winter, the species migrates to the shallow subtidal zone and – although such an impact has not been reported yet in European waters – it could cause damage to natural or cultivated shellfish stocks (Dauvin 2009). Along the western Atlantic coast, it has been observed to reach very high densities and have negative impacts on small recruits and juveniles of several native species of barnacles, littorine snails, brachyuran crabs, and mytilid bivalves (Lohrer and Whitlatch 2002a,b). H. sanguineus competes for food and space with the native crab Carcinus maenas and has been reported to cause its decline in some invaded areas (Dauvin et al. 2009; Landschoff et al. 2013E; Epifanio 2013), while predation of newly settled C. maenas by H. sanguineus has also been documented (Lohrer and Whitlatch 2002aE). The species can serve as an important food resource for birds and fish (Dauvin and Dufossé 2011).

Hemimysis anomala: The bloody-red shrimp is a voracious predator and an opportunistic omnivorous feeder (Ketelaars et al. 1999). H. anomala redirects energy from pelagic to benthic subsystems and provides additional prey to planktivorous and benthivorous fish (Leppäkoski et al. 2002b; Dumont and Muller 2010). Populations of zooplankton (in particular cladocerans) can decrease following the establishment of H. anomala (Ketelaars et al. 1999). However, these impacts have been reported mainly for freshwater systems and not for marine areas. The species is reported to tolerate salinities up to 19‰ (Kelleher et al. 1999); however, so far, the abundant populations were found only in fresh and oligohaline waters. It seems that the species is less invasive in marine brackish waters; no impact on biodiversity or ecosystem services has been reported for marine waters.

Homarus americanus: American lobsters may compete with the native commercially exploited European lobster and possibly Norwegian lobsters, by sharing the same habitat and food preferences (van der Meeren et al. 2010 and references therein). However, the status of the species in Europe is not well understood and it does not seem to be well established in most areas (van der Meeren et al. 2010; Stebbing et al. 2012a). Although hybridization is possible, European lobsters are able to recognize conspecific mates, perhaps by chemical signals, and these signals serve as pre-mating barriers preventing hybridization (van der Meeren et al. 2008, 2010). The European lobster stocks in the Nordic region are vulnerable and the presence of a competitive sibling species could be harmful. American lobsters carry diseases that might have important negative effects on European lobster stocks (van der Meeren et al. 2010). Gaffkaemia, caused by Aerococcus viridans var. homari, has been introduced in European waters


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by American lobsters and has infected European lobsters both in holding facilities and in the wild; in holding facilities, Gaffkaemia has caused severe mortalities of European lobsters (Stebbing et al. 2012b).

Marsupenaeus japonicus: The species entered the Levant Sea through the Suez Canal; since the 1980s it has also been farmed in southern Europe (Savini et al. 2010 and references therein). M. japonicus is of major commercial importance, a highly prized species considered a boon to the Levantine fisheries as it brings in a substantial part of trawl catches and income (Galil 2007a and references therein). However, this came in expense of the native penaeid prawn, Melicertus kerathurus, which was common in the Levantine Sea and supported commercial fisheries throughout the 1950s. M. kerathurus has now almost disappeared and its habitat overrun by M. japonicus (Galil 2007a).

Monocorophium sextonae: There are no documented impacts on ecosystem services or biodiversity in Europe.

Palaemon elegans: P. elegans is known to colonize rapidly inshore, lagoon and estuarine habitats subsequently outcompeting and replacing native palaemonid shrimps such as P. adspersus and Palaemonetes varians (Grabowski 2006; CABI 2013). In the invaded territories, P. elegans is an opportunistic feeder with a broad diet including detritus, algae and, having the highest share, a number of zoobenthic crustaceans, including gammarids (Janas and Barańska 2008). As P. elegans may consume large quantities of gammarid amphipods, an introduction of the alien palaemonid may exert a strong predatory pressure on another invasive species, Gammarus tigrinus (Katajisto et al. 2013). P. elegans forms abundant populations and its energetic value is not less than in the native P. adspersus (Janas and Bruska 2010) so it may enrich the food base of various bird and fish species (Gruszka and Więcaszek 2011; CABI 2013).

Palaemon macrodactylus: P. macrodactylus is an invader of transitional waters. It is an edible prawn that can be exploited by coastal fisheries. There are concerns that it might outcompete native species, such as the commercially exploited Palaemon longirostris, but there is yet no such evidence from European waters (Chícharo et al. 2009; Lavesque et al. 2010). It was, however, reported that P. macrodactylus outcompeted and replaced the native Crangon shrimp (Ricketts et al. 1968) so we cannot exclude the possibility of posing some threat to native shrimp in Europe.

Paralithodes camtschaticus: The very large red king crab is an omnivorous predator and grows up to > 220 mm carapace and a span from leg to leg up to 140 cm; its weight may exceed 10 kg (DAISIE 2013). It is an active predator on benthic fauna, especially feeding in deep soft-bottom environments, and having an enormous predatory impact on local species, especially during mass developments (DAISIE 2013). It can cause reduction of benthic biodiversity and substantial reductions in populations of echinoderms, polychaetes, and bivalves (Britayev et al. 2010; Oug et al. 2011). It can cause degradation of sediment habitat quality due to hypoxic conditions and low biological activity (Oug et al. 2011). The latter is probably caused by the reduction of the functional diversity of the resident species assemblages (by removing organisms performing important functions such as bio-irrigation and sediment reworking), which may have overall implications for ecosystem function, production and responses to other environmental stressors (Oug et al. 2011; Falk-Petersen et al. 2011). Impacts on commercial and non-commercial fish species, through egg predation or competition for prey is possible (Falk-Petersen et al. 2011 and references therein). The species is highly valued and supports a profitable fishery in Norway and Russia; due to the commercial interest, a total annual catch limit was set to avoid over-fishing of the species (DAISIE 2013). P. camtschaticus impacts the longline fishery by removing bait from hooks, thereby reducing catches of targeted fish (Sundet and Hjelset 2002).

Percnon gibbesi: There is no reported impact of this species on ecosystem services or biodiversity. Such impacts on shallow hard-substrate benthic communities and possible cascade effects on trophic webs may not yet be predicted in the absence of information on its grazing intensity and the trophic interactions with other species (Katsanevakis et al. 2011). As the species keeps expanding rapidly and may reach densities of many individuals per m2 (Sciberras and Schembri 2008; Raineri and Savini 2010), its invasion in the shallow rocky infralittoral zone of the Mediterranean Sea may add further stress to the already altered ecosystems, due to overgrazing. P. gibbesi is a potential competitor for space with the native species Pachygrapsus marmoratus (and to a lesser extent with Eriphia verrucosa); however, laboratory experiments indicate that P. marmoratus is unlikely to be excluded from its natural habitat by the alien species and significant spatial resource partitioning on the part of P. marmoratus is unlikely to occur (Sciberras and Schembri 2008).


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Platorchestia platensis: The species has been reported to outcompete the native species Orchestia gammarellus (Karlbrink 1969; Persson 2001). There is no documented impact on ecosystem services.

Portunus segnis (ex Portunus pelagicus): It is an omnivorus species with a preference for crustaceans (Pazooki et al. 2012). It competes for food with native biota (DAISIE 2013) but no displacement or substantial impact at population level has been reported for any native species. The species is commercially important in nearshore fisheries in the Levantine Basin, where it has been marketed since the early 1900s (Munro Fox 1924; Galil 2000).

Rhithropanopeus harrisii: Harris’s mud crab may alter ecosystem functioning by competing with native crabs or even benthophagous fishes, being a predator on native benthic invertebrates, and being a prey of native predators (Roche and Torchin 2007). In the inner parts of the Baltic Sea, where there are no native crabs, this functional species, completely new to the area, has locally dominated zoobenthos (Grabowski et al. 2005). Thus, it has probably impacted the original food web structure. It has been commonly found in the stomachs of several benthophagous fishes, which indicates that top-down regulation through predation by fish may be taking place (Fowler et al. 2013). The crab was identified as a carrier of the white spot syndrome (WSS), a viral infection causing highly lethal and contagious disease in commercially harvested and aquacultured penaeid shrimp (Payen and Bonami 1979). In the Caspian Sea, massive occurrence of R. harrissi created an economic problem by fouling water intake pipes and damaging fishes in gill nets, causing economic loss to local fishermen (Zaitsev and Öztürk 2001). Due to its euryhalinity, the crab colonised almost freshwater reservoirs in Texas and is reported to foul PVC intakes in lakeside homes and clog the cooling system of a nuclear power plant in Glenrose (GISD 2013). On the other hand, R. harrisii has become a popular model organism in many developmental, physiological and ecotoxicity studies (GISD 2013).

Insecta

Telmatogeton japonicus: T. japonicus is a marine chironomid insect, whose larvae live in tubes attached to natural or artificial hard surfaces in the intertidal splash zone. It is common on, and readily takes advantage of the growing presence of, offshore and coastal man-made constructions and artificial substrata, such as breakwaters, concrete beach walls, and offshore windmills and buoys, in the northeastern Atlantic and the Baltic Sea (Wilhelmsson and Malm 2008; Brodin and Anderson 2009; Raunio et al. 2009; Kerckhof et al. 2010). It plays a role as pioneer, starting from zero and building up a flourishing subsystem as dominant species (Kronberg 1988). Consequently, it contributes to both structural and functional diversity of the low-diversity splash zone community. T. japonicus can be advantageous for native species as a food source, especially for birds in autumn and winter when other insects are uncommon and food generally is scarse or less available (Sunose and Fujisawa 1982). Migrating wader birds (Charadriidae) are known to use T. japonicus as an important source of food (Boudewijn and Meijer 2007). Dense tubes may modify microhabitats, but no negative impacts of the species on ecosystem services or biodiversity have been reported (Jensen 2010f).

Mollusca

Anadara kagoshimensis [Bivalvia]: It can become a dominant species on shallow sandy bottoms (Streftaris and Zenetos 2006; Shiganova 2008; Zolotarev and Terentyev 2012), replacing native species, such as the exploited bivalve Cerastoderma glaucum in the N. Adriatic (Occhipinti Ambrogi 2000). It is considered as a habitat engineer. Along the Bulgarian coasts it reaches biomass of up to 4,300 g m-2 but no significant negative impact has been reported (Sahin et al. 2009). Although the species is edible in its native region, attaining high market values, it is not appreciated as food in Italy (Zenetos et al. 2004). However its potential for aquaculture has been investigated in Turkey (Acarli et al. 2012). As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). On the other hand, bioturbation of sediments increases sediment water and oxygen content and releases nutrients from the sediment to the water column (Vaughn and Hakenkamp 2001), thus positively impacting ocean nourishment.


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Anadara transversa (ex A. demiri) [Bivalvia]: A. transversa has become very abundant in many Mediterranean sites, especially in polluted areas (Galil and Zenetos 2002; Çinar et al. 2006b), and has impacted the sandy infralittoral bottoms, negatively affecting populations of commercial species such as the venerid Chamelea gallina (Morello et al. 2004; Zenetos et al. 2010). The density of Anadara transversa was found to be 300 ind m-2 on grey mud and 30 ind m-2 on black mud in Izmir Bay (Aegean Sea) (Demir 1977). It is considered as a habitat engineer; it may reach very high densities in the soft substrata, and the accumulation of empty shells creates a modified environment offering substrate to other organisms (Sousa et al. 2009). A. transversa juveniles may attach to ropes used for mussel culture, negatively affecting mussel settlement (Morello et al. 2004). As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). On the other hand, bioturbation of sediments increases sediment water and oxygen content and releases nutrients from the sediment to the water column (Vaughn and Hakenkamp 2001), thus positively impacting ocean nourishment.

Arcuatula (ex. Musculista) senhousia [Bivalvia]: This mussel is capable of marked habitat alteration (habitat engineer) through the construction of byssal mats on the surface of soft sediments (Crooks 1998). Such mats can alter sedimentary properties, i.e. increase of fine sediment percentages, combustible organic matter percentages, and sediment shear strengths (Crooks 1998). The presence of these mats dramatically alters the resident macroinvertebrate assemblage (CABI 2013). These mats also alter resident macrofaunal assemblages by shifting communities from suspension feeding to primarily deposit-feeding (Crooks 1998; Mistri 2004a; DAISIE 2013). The rate at which large A. senhousia assemblages filter the water for food and oxygen also accelerates the rate of conversion of suspended sediment to deposited material and may rapidly alter the stability of the substrate (CABI 2013). This deposition of large amounts of organic matter alters the nutritional quality of the sediment and leads to the diminution of the redox potential discontinuity layer, rendering the environment within or under byssal mats unsuitable for adults or larvae of other species (Mistri et al. 2004). On the other hand, the creation of a novel three-dimensional episurface structure can provide increased habitat heterogeneity on an otherwise soft bottom and have a positive effect on at least a part of the benthic community (Mistri et al. 2003; Munuary 2008). The presence of A. senhousia mats has been claimed to create heavy impacts on shellfish exploitation of commercially important bivalves through the reduction of shellfish growth and survival, but there is no strong evidence of such an effect (Mistri 2004b).

Brachidontes pharaonis [Bivalvia]: It locally displaces the native mytilid, Mytilaster minimus, by interfering with its recruitment and detrimentally affecting its survival and growth (Safriel and Sasson-Frosting 1988E; Galil 2007a; Crocetta et al. 2013). In the early 1970s it was much rarer than the native M. minimus, that formed dense ‘Mytilaster beds’ but in the late 1990s a rapid shift in dominance was observed, with some Brachidontes populations of up to 300 specimens per 100 cm² on areas where mussel beds were absent in the past, while M. minimus was only rarely encountered (Mienis 2003; Rilov et al. 2004; Galil 2007a). B. pharaonis can deplete the phytoplankton concentration in the water column, constraining the growth of other filter-feeding animals such as Mytilaster minimus. The establishment of massive beds of B. pharaonis and its continuing expansion in the Mediterranean (Sarà et al. 2013) has had significant effects on the biota of intertidal rocky areas, especially on the ecology of the vermetid platforms (Rilov and Galil 2009). The species can also become very abundant in shallow subtidal rocky habitats, reaching depths down to 14m (Crocetta et al. 2013). Being a habitat-forming species, its massive beds have changed the identity and diversity of biocommunities on these platforms by excluding some species and facilitating others. Habitats are clearly impacted when the density of mussels is high, as recorded in western Sicily (375 individuals/400cm2 in saltpans and 10,000 individuals/m2) (Sarà et al. 2006) and in Malta (16,550 individuals/m2) (Bonnici et al. 2012). As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). There was probably a positive impact to the population of the native whelk, Stramonita haemastoma (Linnaeus, 1758), that was found to preferentially prey on Brachidontes (Rilov et al. 2002). The species may induce substantial costs by fouling intake pipes, heat exchangers and underwater constructions (Garaventa et al. 2012; Otero et al. 2013).


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Chama pacifica [Bivalvia]: The spreading of C. pacifica in the Eastern Mediterranean is documented/reviewed by Crocetta and Russo (2012). C. pacifica has formed dense populations in the form of massive oyster beds in many locations of the eastern Mediterranean, at a wide depth range (1–25 m), reaching a density of 13.6 ± 5.3 m-2 at 5 m (Zurel et al. 2011). It is a habitat engineer at high densities, increasing spatial complexity of the benthic habitat. The surface of its shell provides strongholds for a diverse community of algae and invertebrates (Fishelson 2000). C. pacifica reefs provide services similar to oyster beds, such as water quality regulation and bioremediation of waste by biofiltration, climate regulation (by acting as a carbon sink), and supporting reproduction and nursery areas for a variety of species. It is reported to have almost completely replaced the native Chama gryphoides and at some sites the thorny oyster Spondylus gaederopus (Galil 2007b; Otero et al. 2013; Crocetta et al. 2013). Competition and reduced plankton availability caused by decreased water flow can also slow down the growth of other benthic organisms (Otero et al. 2013). As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). It is a valuable species for seashell collectors, with a small trading market (Otero et al. 2013).

Crassostrea gigas [Bivalvia]: The Pacific oysterwas introduced to Europe for aquaculture and soon established wild populations and expanded rapidly, forming extensive and dense reef structures. The introduction of C. gigas has had a highly significant economic impact as a farmed product, and it has become the most important species in oyster culture, as it has great fecundity, is highly resistant to pathogens and diseases and grows to marketable size more quickly than the native O. edulis (Otero et al. 2013). In several countries, the introduction of C. gigas has resulted in building a sustainable shellfish industry providing direct revenues for thousands of farmers and concomitant activities (CABI 2013). A highly valuable indirect economic impact concerns the lasting establishment of coastal communities in otherwise unfavourable rural areas, therefore playing a significant role in coastal management values (CABI 2013). Pacific oysters directly introduced from the wild have been a source of several cryptic diseases, invasive seaweeds, oyster pests and other alien species (DAISIE 2013). Extensive settlements can lead to competition with native biota for both food and space. There are some concerns for mussel cultivation due to heavy settlement and fast growth, with competition for food and space (DAISIE 2013). C. gigas induces changes in plankton composition, habitat heterogeneity and biodiversity, carrying capacity, food webs and parasite life cycles (Troost 2010). Massive increases in C. gigas abundances have led in some cases to the overgrowth of native mussel beds, turning them into oyster reefs (Diederich 2006; Kochmann et al. 2008E). A shift in dominance from mussels to oysters alters habitat structures which entail differential abundances of associated organisms (Kochmann et al. 2008E). In a field experiment, biogenic structures of the mussels and C. gigas, alone or in combination, differentially altered sediment composition, abundance of an oligochaete and polychaete species, recruitment of oysters and immigration of juvenile shore crabs and of periwinkles (Kochmann et al. 2008E). In a natural experiment, Markert et al. (2009)N found increased species richness, abundance, biomass, and diversity in C. gigas reefs in comparison to M. edulis reefs. Mussel beds generally constitute hot spots with respect to productivity and filtering-capacity, biodiversity, and as a food resource for various crustaceans, fish, birds and man, and thus their overgrowth or possible displacement by oysters might profoundly change the entire ecosystem processes and affect all ecosystem services provided by mussel beds, i.e. food provision, recreation, symbolic and aesthetic values, climate regulation, and life cycle maintenance (Salomidi et al. 2012). On the other hand, C. gigas is an ecosystem engineer creating biogenic reefs and providing similar services to mussel beds (Troost 2010). Mature C. gigas reefs appear more persistent than mussel beds and may therefore stabilize the sediments on a larger time scale. These mature reefs are anchored deep in the sediment, thereby consolidating the substrate firmly and thus preventing erosion of intertidal flats locally protecting the intertidal habitat of native bivalves and other invertebrate fauna, and the intertidal foraging grounds of species at higher trophic levels such as shorebirds (Troost 2010). C. gigas reefs may reintroduce structural complexity and may be considered to restore habitat diversity and biodiversity in degraded mussel beds. C. gigas reefs affect consumer-resource dynamics far beyond their own boundaries affecting the distribution of avian predators (van der Zee et al. 2012). As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such


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increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). Furthermore, mature oyster reefs are usually avoided by fishermen because they can cause damage to netting and hence the associated benthic community may be given better opportunities to mature in the more stable environment of an oyster reef, compared to native biogenic structures (Troost 2010). On the other hand, in the Sylt-Rømø Bight ecosystem (North Sea), Baird et al. (2012), based on ecological network analysis (ENA), reported a decline in the number of cycles, trophic efficiency, ascendency ratios, and relative redundancy that was ascribed to the impact of the invasive species C. gigas and Austrominius modestus on system organization and function. Natural hybridization between genetically differentiated populations of C. gigas and C. angulata was demonstrated (Huvet et al. 2004). Therefore, the remaining populations of the C. angulata ecotype in Portugal are at threat from current culture development and extensive transfers of C. gigas (Huvet et al. 2000).

Crassostrea virginica [Bivalvia]: The eastern or American oyster was introduced in many European countries for aquaculture but in most cases the species either did not get established in the wild and is no longer present, or its status is unknown (Minchin 2007; Skolka and Preda 2010; Minchin et al. 2013).

Crepidula fornicata [Gastropoda]: The American slippershell species may occur at densities exceeding 1700 m-2 and attain a wet biomass of 10 kg m-2 (DAISIE 2013). C. fornicata has the potential to act as significant mortality factor as well as a cause of reduced growth rates to Mytilus edulis populations, mainly due to interference and to a lesser extent due to trophic competition (Thieltges 2005aE). At the same time it provides protection to mussels against starfish predation (Thieltges 2005bE). Another positive effect of C. fornicata, especially on the coast of France where it is superabundant, may be that it causes a shift of phytoplankton blooms from toxic flagellates to diatoms (Thieltges et al. 2006). Although initially called an “oyster pest”, no effect of C. fornicata has been found on the survival and growth of oysters (de Montaudouin et al. 1999E), but there is strong competition for space in some areas causing serious problems in shellfish production (Frésard and Boncoeur 2006; Blanchard 2009). Dense limpet populations disturb fishery or oyster farming activities to such an extent that in some bays in Great Britain and in France, cleaning operations are necessary (CABI 2013). An additional cost for shellfish production is for sorting and cleaning shells fouled by C. fornicata before marketing (Blanchard 1997). Recently, exploitation of the species for human consumption has started in France (http://www.

nytimes.com/2014/03/12/world/europe/in-france-a-quest-to-convert-a-sea-snail-plague-into-a-culinary-pleasure.html?hp&_r=2). This species is a habitat engineer and has been reported to cause substantial large-scale changes on the recipient ecosystems such as modification in the trophic structure, changes in phytoplankton composition, enhanced siltation due to accumulation of faeces and pseudofaeces, and changes in benthic sediments and near-bottom currents (Thieltges et al. 2006 and references therein). Such accumulating sediments on maerl beds cause their degradation (DAISIE 2013) and negatively affect all related ecosystem services such as provision of food and biotic materials, climate regulation, cognitive benefits, primary production, support of reproduction and nursery areas (Salomidi et al. 2012). Dense populations spread and completely cover the ground, so that the sediment disappears under the stacks and has no more exchange with the water. By trapping the suspended matter and producing lots of mucous pseudofaeces, a dense population transforms the primary sandy sediment into a muddy one with a high organic content, which becomes rapidly anoxic and unsuitable for other species (CABI 2013). The original endofauna often disappears completely when new epifauna increases on the stacks and in the interstices. A new limpet community appears, with a lot of suspension-feeders (bryozoans, ascidians, fixed annelids and cirripeds) and several little carnivorous species of crustaceans in the interstices (CABI 2013). The species may also reduce recruitment of some benthic commercial fishes, e.g. a negative role of C. fornicata on juvenile sole density with possible effects on nursery habitat capacity and sole stock recruitment has been reported (Le Pape et al. 2004). It also fouls underwater constructions and artificial structures in ports (DAISIE 2013). On the other hand, C. fornicata has some important positive impacts on many species or ecosystems: it reduces predation pressure to basibionts, provides additional substrate for other epibenthic species, adds heterogeneity to habitat structure, reduces parasite attacks on basibionts, may improve water quality and reduce toxic algal blooms, and may increase diversity, biomass and abundance of zoobenthic communities (de Montaudouin and Sauriau 1999N; Ragueneau et al. 2002N; Prinz 2005E; Thieltges et al. 2006; Thieltges et al. 2009E). C. fornicata has been dredged on a large scale and used as a calcareous supplement for agriculture (Blanchard 2009).


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Ensis directus (ex Ensis americanus) [Bivalvia]: The American razor clam E. directus has become a prominent component of the macrobenthos in shallow subtidal sands of the North Sea; in many areas it is the most abundant large bivalve in the shallow subtidal. However, there is no evidence that E. directus suppresses populations of native species. On the contrary, E. directus seems to favour the settlement of some deposit feeders (Dannheim and Rumohr 2012). The species is edible and is commercially exploited; it has now become a very common bait used for surfcasting in Italy (Crocetta, pers. comm.). Dense clam mats might stabilise the sediment and function as a sediment-trap for organic matter. As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). On the other hand, bioturbation of sediments increases sediment water and oxygen content and releases nutrients from the sediment to the water column (Vaughn and Hakenkamp 2001), thus positively impacting ocean nourishment. In many areas, E. directus has become an important food item for fish and seabirds (Tulp et al. 2010). The species currently plays an important role for the higher trophic levels and its introduction has caused a major change in food relations in its distribution area (Tulp et al. 2010). Diet studies showed that E. directus makes up a significant proportion (20–100% of the total weight in fish stomachs) in the current diet of plaice (Pleuronectes platessa), sole (Solea solea), dab (Limanda limanda), flounder (Platichthys flesus) and dragonet (Callionymus lyra) and in the diet of common eiders (Somateria mollissima) (85-90% occurence) and common scoter (Melanitta nigra) (26% occurrence); waders, gulls and corvids also prey on E. directus (Tulp et al. 2010). E. directus might have an impact on the recreational value of some beaches as its sharp shells can cause deep cuts on bathers’ feet. Such injuries can also occur when stepping on native species, but E. directus lives at much shallower depths than native species and consequently injuries are more likely (DAISIE 2013). Mechanical removal of dead shells is occasionally carried out (CABI 2013).

Mercenaria mercenaria [Bivalvia]: The northern Quahog is a much-esteemed edible bivalve has been introduced several times for aquaculture in many European countries. It is possibly important in pharmacology, as the visceral mass and especially the liver and the crystalline style, contain a substance capable of acting with good selectivity on cancerous cells (Schmeer 1979). Wild populations are now present in some locations, e.g. in Brittany. In Europe, there are no documented negative impacts on ecosystem services or biodiversity (Jensen 2010c).

Mya arenaria [Bivalvia]: The soft-shell calm M. arenaria is considered completely naturalized in the Northern seas, as it has been present for more than 500 years, and thus it is difficult to identify its impacts (Jensen 2010d). However, in relatively newly invaded areas (e.g. in the Black Sea) it shows invasive properties dominating in the soft substratum communities, causing regime shifts and replacing native species, causing structural changes in native communities, affecting sediment and water-column characteristics and modifying invaded habitats (Hansen et al. 1996N; Olenin and Leppäkoski 1999; Riisgard and Seerup 2003; Gomoiu 2005; Shiganova 2008; Skolka and Preda 2010). It is a new invader in the Mediterranean Sea (Crocetta and Turolla 2011; Crocetta 2012), and no impacts have been reported therein so far. In the Black Sea, Mya is a key species of a biocoenosis covering about 1000 km2 of the northwestern Black Sea shelf. It has replaced native dominant species such as the small bivalve Lentidium mediterraneum and caused a noticeable impact on the benthic community structure and its biodiversity, as the total number of invertebrate species in the Mya biocoenosis is 2.5 times lower than in the original Lentidium community (Zaitsev and Öztürk 2001). In the Baltic Sea, there seems to be some interaction causing the native clam Macoma balthica to decrease in biomass in places where M. arenaria is abundant (Obolewski and Piesik 2005). M. arenaria’s high abundance, high filtration capacity, ecosystem engineering characteristics and importance in food-web interactions, suggest that this species has dramatically impacted shallow coastal ecosystems not only in Denmark, but potentially throughout invaded locations elsewhere in Europe (Thomsen et al. 2009). Mya arenaria has a high capacity for filtration (Jørgensen and Riisgård 1988; Foster and Zettler 2004) and is able to substantially reduce phytoplankton concentration (Gomoiu 2005; Petersen et al. 2008). It is a food source for many fish (such as gobies, flounder, turbot and sturgeons) and sea birds, and it is also used in some localities to feed poultry (Gomoiu 2005; Shiganova 2008). M. arenaria is an ecosystem engineer. It seems often to die in situ, forming so-called death assemblages (Strasser 1999), which can persist for many decades forming a novel habitat (Palacios et al. 2000). Surfacing shells provide refuge to juvenile crabs and other


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invertebrates and also affect hydrodynamics (Sousa et al. 2009). It sequesters carbon in the form of calcium carbonate used for shell creation. As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). On the other hand, bioturbation of sediments increases sediment water and oxygen content and releases nutrients from the sediment to the water column (Vaughn and Hakenkamp 2001), thus positively impacting water nourishment. Live clams likely enhance oxygen and solute penetration to deep sediment layers because of deep burrowing and play an important role in production, mineralization and redistribution of organic matter in intertidal environments (Sousa et al. 2009). Because of its size (to 10 cm), abundance, longevity (max. 28 years) and ease of identification, M. arenaria is used as a biomonitor and indicator species in several Baltic countries (Jensen 2010d). In its native area M. arenaria is a highly valued food item.

Mytilus edulis [Bivalvia]: The blue mussel is considered to be alien in the Black Sea, where it may hybridize with M. galloprovincialis (Alexandrov et al. 2007). No impacts on biodiversity and ecosystem services have been reported.

Petricolaria pholadiformis [Bivalvia]: It has been reported that P. pholadiformis can displace native species such as Barnea candida (Eno et al. 1997) and Pholas dactylus, but these records seem rather anecdotal and unverified (Jensen 2010e). P. pholadiformis bores into clay, peat, mud, sand and other soft sediments, and may therefore modify habitats (CABI 2013). Its burrowing creates a generally uneven surface on a small scale (5-15 cm) providing habitats for other animals that inhabit vacant burrows and crevices in the clay (Marshall 2008).

Pinctada imbricata radiata [Bivalvia]: This thermophilic bivalve is very abundant in the eastern and southern Mediterranean, and at some sites it can reach densities of many hundreds of individuals per square meter. It is considered a habitat-modifying, gregarious bivalve capable of impacting native fauna by forming oyster banks (DAISIE 2013). It fouls mussel lines and commercial shellfish collectors, negatively impacting shellfish farming activities (DAISIE 2013).The species is edible and has been introduced for mariculture in many areas of Greece and Italy during the last century (Katsanevakis et al. 2008). The introduced oyster may play the role of a habitat engineer at high densities, increasing spatial complexity of the benthic habitat and influencing the trophic pattern of its fauna (Tlig-Zouari et al. 2011). Pinctada imbricata radiata reefs provide services similar to mussel and oyster beds, such as food provision, water purification by biofiltration, and life cycle maintenance for a variety of species. As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). Because of its tolerance to chemical contamination and its ability to accumulate metals to several orders of magnitude higher than the background medium, it has been used as a bioindicator of heavy metal pollution (Göksu et al. 2005).

Potamopyrgus antipodarum [Gastropoda]: The euryhaline mud snail P. antipodarum, although mostly thriving in freshwater habitats, is also found in brackish and even salty waters up to 26 PSU. It is known to disperse by both passive (birds, fish, drift, floating macrophytes) and active (positive rheotactic) mechanisms. The New Zealand mud snail has invaded estuarine areas of all European regional seas. Higher impacts have been reported in freshwater systems than in marine and estuarine areas (Alonso and Castro-Díez 2012). This successful early colonizer is tolerant of a wide range of environmental conditions and has a very high parthenogenetic reproductive capacity, which may lead to the establishment of very dense populations of thousands ind m2 [several authors reported densities up to 500,000 ind m2 in invaded habitats or even up to 800,000 ind m2 (Alonso and Castro-Díez 2012 and references therein)], a fast spread, a high consumption rate of the available primary production of the impacted ecosystem, and domination in the incipient community (Alonso and Castro-Díez 2008). P. antipodarum may alter ecosystem dynamics, compete with and displace native invertebrates, and negatively influence higher trophic levels. In ditches and canals of the basin of the Mont St-Michel Bay (France), this mud snail dominates the gastropod communities in fresh- and salt-water ecosystems (Gérard et al. 2003). Fish and birds are known to ingest mudsnails which, however, may pass the digestive canal alive: thus the energetic benefit to the fish might be low (Aarnio and Bonsdorff 1997;


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Alonso and Castro-Díez 2012). Other reported impacts of the species (on nutrient cycles, native fauna, fish health) are mostly relevant to freshwater bodies and not to European marine and coastal ecosystems.

Rapana venosa [Gastropoda]: The veined rapa whelk is considered a serious menace to bivalve fisheries (Savini and Occhipinti-Ambrogi 2006 and references therein), feeding mainly on mussels, oysters, clams and other bivalves. It has been responsible for the depletion of large stocks of commercial bivalves (esp. Mytilus galloprovincialis) and the associated communities in the Black Sea since the 1950s (Zolotarev 1996; Salomidi et al. 2012). Its distribution is associated with reduction in the range and density of oyster and mussel settlements, in particular near the coasts of Anatolia and Caucasus. In Ukrainian waters, it destroyed the oyster banks in the area of the Kerch Strait and in Karkinitsky Bay. Hence, the species has a severe impact on all ecosystem services provided by mussel and oyster biogenic reefs, i.e. food provision, water purification, coastal protection, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012). R. venosa may become a competitor of the native whelks Buccinum undatum in the North Sea and Stramonita haemastoma in the Mediterranean. The species is edible and has supported profitable fisheries in the Black Sea (Çinar et al. 2005). For the first time in 1982, this species gained an economic importance and was exported as Rapana meat to Japan. It is also locally consumed by the Black Sea residents. The fishery of R. venosa was intense along the Turkish Black Sea coast, reaching up to 1166 t in 1986, yielding 3,415,884 US $ (Bilecik 1990), and has increased in recent years. The annual R. venosa catches from Turkey and Bulgaria reached a total of 13,000 t in 2005 (Sahin et al. 2009, as R. thomasiana). However, in the easternmost Black Sea, R. venosa fisheries ceased operating from 2005 until recently due to a diminishing mean size of the animal (Ulman et al. 2013). Empty shells may be marketed as tourist souvenirs (Otero et al. 2013). Illegal bottom trawling for harvesting of R. venosa along the Black Sea shelf has raised ecological concerns with respect to the benthic communities and especially the mussel beds (Knudsen et al. 2010; Ulman et al. 2013). It utilises nets as spawning substratum, adding extra load to the draught (CABI 2013).

Spondylus spinosus [Bivalvia]: Within a decade of its first record in Haifa Bay (Israel), S. spinosus has been reported to completely outcompete the native congener S. gaederopus along the eastern Mediterranean shores (Mienis et al. 1993; Crocetta et al. 2013). S. spinosus can thrive in harbour environments and create dense aggregations, generating solid reefs, and thus acting as an ecosystem engineer (Streftaris and Zenetos 2006). It forms dense and strong populations along with Chama pacifica and other Spondylidae, the surfaces of their shells providing strongholds for a diverse community of algae and invertebrates (Fishelson 2000). It sequesters carbon in the form of calcium carbonate used for shell creation. Competition and reduced plankton availability caused by decreased water flow can also slow down the growth of other benthic organisms (Otero et al. 2013). As all filter feeders, it filters suspended particles and subsequently deposits faeces as well as particles captured but not consumed onto the sediments, increasing sedimentation. Such increased sedimentation can represent a significant loss of energy and nutrients from the water column and decrease pelagic production (Strayer et al. 1999; Vanni 2002; Newell 2004). The economic impact of this invasive species is unknown. Some members of the family are known for their decorative shells and have been adequately studied from the perspective of aquaculture (Zenetos et al. 2004). It is served in seafood restaurants in Israel and Lebanon (Rilov and Galil 2009).

Teredo navalis [Bivalvia]: This is a cryptogenic (Hoppe 2002) wood-boring marine bivalve (‘shipworm’) that first appeared in western Europe in 1730 and within a couple of years caused enormous damage to wooden dyke gates in the Netherlands (Paalvast and van der Velde 2011). Recently, damage caused by the species along the German coast in the western Baltic was estimated to amount to approximately 25 - 50 million Euros (DAISIE 2013). It destroys archaeologically valuable ancient wooden shipwrecks (Förster 2003). The species occupies an exclusivist ecological niche and does not compete with any native species.

Urosalpinx cinerea [Gastropoda]: The Atlantic oyster drill (Urosalpinx cinerea) prays on oysters, mussels and clams and may compete with native gastropods. It is a major pest to the commercial oyster industry. It is especially harmful to oyster spat and in the past it inflicted a ~50% mortality on Essex oyster beds (UK) before its populations declined in the 1980s (Hancock 1954).

Venerupis (ex Ruditapes) philippinarum [Bivalvia]: The Manila clam, V. philippinarum, was intentionally introduced to many European coastal areas, aiming at compensating for irregular yields of the native European sister species Ruditapes decussatus, one of the main shellfish industries in Western


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Europe, and enhance the depressed fisheries and aquaculture activities. V. philippinarum currently supports local fisheries in many European coastal areas and may reach densities over 1000 specimens per square meter, dominating macrobenthic communities in terms of abundance and biomass. It is one of the most important species in shellfish farming and its production accounts for 20% of the global shellfish market (Otero et al. 2013). Italy is the largest European producer of V. philippinarum with 90% of the European market, worth over 100 million euros (Otero et al. 2013). It is a marine suspension feeding bivalve and an important ecosystem engineer known to significantly increase sediment erosion and re-suspension rates and reduce sediment stability (Sgro et al. 2005E); evidence of significant effects on community bioturbation rates and sediment mixing has been reported, influencing the distribution of oxygen in marine sediments, enhancing solute exchange with the overlying water column, and affecting nutrient cycling (Bartoli et al. 2001N; Queirós et al. 2011). The rapid circulation of nutrients due to R. philippinarum dense populations can promote new phytoplankon blooms and also sustain macroalgal growth (Bartoli et al. 2001N), and thus positively affect primary production. In the Venice lagoon, it was estimated that with the introduction of V. philippinarum the filtration capacity has more than doubled; this has altered the functioning of the ecosystem, resulting in a stronger benthic–pelagic coupling (Pranovi et al. 2006). The latter authors stated that with the introduction of V. philippinarum “the Venice Lagoon ecosystem has entered into a new state, probably more resistant but less resilient”. The culture of Manila clam was responsible for the introduction of a large number of non-native invertebrates and algae, often attached to packaging material, fouling the shell or parasitizing bivalve tissues (Savini et al. 2010). In the Venice lagoon, the spread of V. philippinarum was related to a sharp reduction, both in terms of distribution area and density, of all other filter feeder bivalves (Pranovi et al. 2006). The possible genetic impact of V. philippinarum on the native Venerupis decussata through hybridisation and introgression, threatening populations of the latter, has been reported (Hurtado et al. 2011). On the other hand, the introduction of V. philippinarum into European coastal waters has presented the Eurasian oystercatcher Haematopus ostralegus ostralegus with a new food resource and reduced its predicted over-winter mortality; similar benefits for other shellfish-eating shorebird populations seem probable (Caldow et al. 2007).

Ascidiacea

Botrylloides violaceus: This is an invasive colonial ascidian, commonly recognized as a biofouling nuisance species. In North America, B. violaceus impacts both aquaculture facilities and natural ecosystems (Bock et al. 2011 and references therein). B. violaceus can be a pest to shellfish culture as it overgrows shellfish (e.g. mussels) and other sessile invertebrate species (CABI 2013). However, such impacts on aquaculture have not been reported in Europe. In European waters, it has been reported to overgrow large surface areas and outcompete many of the native species (Gittenberger et al. 2010). Under certain conditions, it may outcompete the native B. schlosseri for space (Gittenberger and Moons 2011N).

Didemnum vexillum: In other (non-European) regions, where D. vexillum is invasive, it had a severe impact on the native ecosystems by overgrowing large areas of the bottom, suffocating virtually every organism (Gittenberger et al. 2010). It also has significant negative impacts on cultured or commercially important wild shellfish. However, in European waters the species is not yet common and there are no documented impacts on ecosystem services or biodiversity.

Diplosoma listerianum: There are no documented impacts on ecosystem services or biodiversity in Europe.

Microcosmus squamiger: M. squamiger has the ability to occupy extensive areas of hard substrata, especially in harbours, bays, and shallow littoral habitats, and to outcompete native species, forming dense aggregations from about 500 to up to 2,300 individuals per m2 (Turon et al. 2007; Mastrototaro and Dappiano 2008; Rius et al. 2009; Otero et al. 2013). It is considered a pest to bivalve culture in some areas, where it competes for food and space (Otero et al. 2013). It is also a nuisance fouler on ships, recreational vessels and other submerged manmade structures (Otero et al. 2013). The gastropod Stramonita haemastoma preys on M. squamiger and a significant positive correlation of T. haemastoma abundance and M. squamiger biomas has been found (Rius et al. 2009).


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Molgula manhattensis: It is a fouling species. There are no documented impacts on ecosystem services or biodiversity in Europe.

Polyandrocarpa zorritensis: There are no documented impacts on ecosystem services or biodiversity in Europe. P. zorritensis may have a negative impact on artificial structures present in shallow waters. The species is not invasive in Europe (CABI 2013).

Styela clava: The club tunicate can attain densities >1000 m-2 in sheltered areas, creating a high biomass that results in competition with other filter-feeders (DAISIE 2013; CABI 2013). In some areas the increase of S. clava populations has been paralleled by a decline in the population of the native ascidian Ciona intestinalis (Lützen 1999). It can foul aquaculture gear and farmed shellfish causing substantial damages and costs (Karney and Rhee 2009; Davis and Davis 2009). In southern Korea, Styela clava is considered to be a seafood delicacy and has acquired a cultural distinction as an aphrodisiac (Karney and Rhee 2009). A consumer market already exists for the species in Korean markets in the U.S. that sell imported frozen S. clava (Karney and Rhee 2009); hence there is a potential for exploring the species in European waters.

Bryozoa

Bugula neritina: It is a fouling pest. There are no documented negative impacts on ecosystem services or biodiversity. Bugula neritina colonies are the source of a novel chemical compound (bryostatin), which has been shown to be effective against leukaemia and a number of other kinds of cancer (CABI 2013). A newly described species of bacterium, which is symbiotic to B. neritina cryptic species 'type D', appears to be the source of bryostatins (Davidson and Haygood 1999; Davidson et al. 2001).

Tricellaria inopinata: This is a fast-growing fouling organism, settling on natural and artificial hard substrata. The invasion of the species in the Venice Lagoon seems to have caused a drastic reduction in frequency and abundance of native bryozoan species, such as Bugula stolonifera and Bugula neritina (CABI 2013).

Victorella pavida: It is known to (re)colonize free space rapidly. V. pavida contributes to species diversity in shallow water habitats by providing habitat for a wide variety of small motile animals. On the other hand, colonies are presumed to compete for space with other fouling taxa, e.g., barnacles and hydroids, and depress them (Carter and Jackson 2007).

Cnidaria

Blackfordia virginica: There are no documented impacts on ecosystem services or biodiversity in Europe (Graham and Bayha 2007).

Cordylophora caspia: In some sites in the Baltic Sea, the Ponto-Caspian invasive species C. caspia can be very abundant and may cause economic damage to fisheries, shipping, boating, fish farming and industry (Leppakoski et al. 2002). It also clogs water intakes for cooling water (Jansson 1994). It can modify the habitat substrate by trapping particulate organic matter in the interstitial microhabitats created by the thick hydroid colonies (Leppakoski et al. 2002). It competes with native species for space and food (DAISIE 2013). This species forms a sparse population in the Mediterranean Sea (Çinar et al. 2008). Large and dense colonies of the hydroids essentially modify benthic habitats causing structural changes in pelagic and benthic communities (DAISIE 2013).

Gonionemus vertens: Stings from G. vertens are unusually venomous. In an area around Vladivostok (Russia) it causes a severe envenomation syndrome at certain times of year (Yakovlev Vaskovsky 1993). However, in Europe there are no documented impacts on ecosystem services or biodiversity (CABI 2013).

Oculina patagonica: This scleractinian coral is very invasive in some localities, especially on anthropogenic hard substrata in disturbed coastal habitats (Sartoretto et al. 2008; Zenetos et al. 2009; Coma et al. 2011; Salomidi et al. 2013; Serrano et al. 2013; Rubio-Portillo et al. 2014). It also occurs in natural substrate in unpolluted waters (Çinar et al. 2006a; Salomidi et al. 2013). It is an "opportunistic dominant settler", competing for space with sessile species, overgrowing many sessile invertebrates, and eliminating algae at its growing edge (Sartoretto et al. 2008; Coma et al. 2011). It out-competes the


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indigenous coral Cladocora caespitosa which is overgrown when the two species come into contact (Sartoretto et al. 2008). O. patagonica is able to initiate a substantial change in community structure and end the monopolization of algae in shallow assemblages; potentially it can modify both the underwater seascape and the sources of primary production in the ecosystem (Coma et al. 2011; Serrano et al. 2012). Communities of sublittoral algae are affected by O. patagonica, which diminishes the structural complexity and richness of species and impacts ecosystem services provided by these biotopes, such as food, biotic materials, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012).

Phyllorhiza punctata: This species has invaded many marine regions worldwide; blooms of P. punctata have been reported to seriously impact fisheries, fish populations and the wider ecosystem functioning in the Gulf of Mexico (Graham et al. 2003; Graham and Bayha 2007). Juveniles of the Lessepsian fish Alepes djedaba were recorded among tentacles of P. punctata, (Çevik et al. 2011). At peak densities (>1 ind/m3), this species was an extensive consumer of fish eggs and zooplankton (Graham et al. 2003; Johnson et al. 2005). This species was reported to be responsible for the unusually low shrimp harvest along the coasts of Mississippi and Louisiana (Graham et al. 2003). However, in European waters, the species rarely occurred in the eastern (Galil et al. 1990) and western (Boero et al. 2009) Mediterranean Sea, and no impacts on ecosystem services or biodiversity have been reported.

Rhopilema nomadica: Since the mid-1980s, massive swarms of this planktivorus jellyfish have appeared along the Levant coast, stretching for more than 100 km (Rilov and Galil 2009). These swarms frequently draw nearer to the shore and adversely affect tourism, fisheries and coastal installations. Local municipalities report a decrease in vacationers frequenting the beaches because of the public’s concern over the painful stings inflicted by the jellyfish (Yoffe and Baruchin 2004). This species inflicts painful stings on swimmers and fishermen, characterized by erythematous eruptions, itching and burning sensations, and its symptoms include fever, fatigue and muscular aches (Galil et al. 1990; Gusmani et al. 1997). In summer 2009, several blooms were observed along the coastline between Antalya and Adana (Turkish Levantine coast) where 815 hospitalized cases were recorded (Öztürk and Isinibilir 2010; Çinar et al. 2011). Coastal trawling and purse-seine fishing are disrupted for the duration of the swarming due to net clogging and inability to sort yield, due to the overwhelming presence of these venomous medusas in the nets (Rilov and Galil 2009). Jellyfish-blocked water intake pipes pose a threat to engine cooling systems of ships and coastal power plants. This species shelters among its tentacles the juveniles of the Lessepsian fish Alepes djedaba and may have caused the sudden population increase of this commercially important fishery (Avian et al. 1995). The proliferation of this species in the eastern Mediterranean resulted in profound changes in the native fauna, such as the replacement of the other large-sized medusa Rhizostoma pulma (Galil 2000). This species was reported to be responsible for veliger mortality and reduced recruitment of Stramonita haemastoma (Rilov et al. 2001).

Ctenophora

Beroe ovata: B. ovata is a predator of the high-impact invasive ctenophore Mnemiopsis leidyi and has been recorded to cause its decline in the Black Sea (Shiganova et al. 2001a; Finenko et al. 2003). Hence, there is a positive impact of B. ovata to those ecosystem services and biodiversity components negatively affected by M. leidyi (see relevant description of M. leidyi impact). The density and biomass of mesozooplankton and its species diversity greatly increased in the Black Sea after the appearance of B. ovata, and were similar to values observed prior to the M. leidyi invasion (Shiganova et al. 2001a). Its appearance led to a partial recovery of the planktonic food web structure in the Black Sea. B. ovata seems not to have any direct negative impact to mesozooplankton or endemic gelatinous species (Aurelia aurita, Pleurobrachia pileus) (Shiganova et al. 2001a).

Mnemiopsis leidyi: The carnivorous ctenophore M. leidyi has invaded the Black Sea, the Mediterranean, the Caspian Sea, the North Sea, and the Baltic. In the Black Sea, the introduction and outbreak of M. leidyi caused dramatic reductions in zooplankton, ichthyoplankton, and zooplanktivorous fish populations in the 1980s and early 1990s (Tsikhon-Lukanina et al. 1993; Shiganova 1998; Shiganova et al. 2001b). This species affected stocks of many commercial fish, especially anchovy Engraulis encrasicolus, Mediterranean horse mackerel Trachurus mediterraneus, and Azov kilka Clupeonella cultriventris (Shiganova et al. 2001b). The annual loss of the fish catch attributed to the


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Mnemiopsis plague was calculated to be approximately 200 million USD in the Black Sea and 30-40 million USD in the Sea of Azov (GESAMP 1997). The population outbreak of M. leidyi in the Black Sea was regarded as one of the most dramatic gelatinous invasion events with profound implications for ecosystem functioning (Kideys 2002). In the Caspian Sea density and biomass of zooplankton decreased substantially by M. leidyi predation as well as catches of three species of kilka (DAISIE 2013). In the Caspian Sea, it was estimated that during winter/spring the population of M. leidyi consumed the available stock of zooplankton in 3 to 8 days, whereas in summer consumption took only 1 day (Finenko et al. 2006). M. leidyi invasion caused cascading effects in the Black and Caspian Seas. The bottom-up effects include the collapse of planktivorous fish, vanishing dolphins in the Black Sea and seals in the Caspian Sea. Top-down effects include an increase in phytoplankton, free from grazing pressure, and increasing bacterioplankton populations, triggering increases in zooflagellates and infusoria populations (DAISIE 2013). Significant economic losses occurred in the Black Sea and Caspian Sea coastal countries due to drastic decline in pelagic fish catches. M. leidyi can cause anoxia in bottom-near waters due to massive deposition of dead individuals (Streftaris and Zenetos 2006).

Echinodermata

Acanthaster planci: It has a high impact in tropical coral reefs. In Europe its presence is questionable (Mediterranean Sea; Zenetos et al. 2010) and there is no documented impact on ecosystem services or biodiversity.

Fish

Liza haematocheila: The so-iny mullet was introduced into the Azov and Black Seas in the 1970s, as a candidate species for aquaculture and fishery enhancement. It has established exploitable populations in the Black Sea and the northern Aegean Sea (Corsini-Foka and Economidis 2007). Along the shores of the Black Sea, its expansion corresponds to a sharp decline of native species of Mugilidae, with which it could compete for food (Erdogan et al. 2010).

Fistularia commersonii: The bluespotted cornetfish is a high-order carnivore preying on native commercially important fish, such as Spicara smaris, Boops boops, and Mullus spp. (Kalogirou et al. 2007; Bariche et al. 2009). It expands rapidly (Azzuro et al. 2013) and there are concerns of possible impacts on the structure and population dynamics of native communities (Kalogirou et al. 2007). F. commersonii is of minor commercial importance; however, it is increasingly acquiring economic significance in eastern Mediterranean local markets, as it has white, palatable flesh and no spines (Otero et al. 2013).

Lagocephalus sceleratus: L. sceleratus, despite being a relatively new invader (first record in the Mediterranean Sea was in 2003 by Filiz and Er 2004), has become abundant and well-established in the Levantine basin (Katsanevakis et al. 2009; Kalogirou 2011). In many areas, it has a significant negative impact on the artisanal fisheries, since it often damages both the fishing gear and the catch of the fishermen, who have to alter their fishing practices (gear, depths, time of the day, etc.) in order to avoid the species (Katsanevakis et al. 2009). L. sceleratus also presents a potential risk to humans, since it contains tetrodotoxin, which may cause poisoning and even death (Bentur et al. 2008; Eisenman et al. 2008; Katikou et al. 2009). Since its arrival in the Mediterranean Sea, it has caused numerous hospitalizations after human consuption; there has been at least one fatality. L. sceleratus preys on various species of fish and invertebrates, some of commercial importance (e.g. Sepia officinalis and Octopus vulgaris) and could potentially affect their stocks (Kalogirou 2011; 2013).

Neogobius melanostomus (synonym: Apollonia melanostoma): The round goby has become dominant in shallow waters of many invaded areas in the southern Baltic Sea, especially in the Gulf of Gdańsk (Sapota 2006). N. melanostomus successfully competes for space, nesting sites and food resources with many other cohabiting benthic fish, such as the flounder, Platichthys flesus, other Gobiidae (especially Pomatoschistus microps, P. minutus, Gobius niger) and the eelpout (Zoarces viviparus) (Corkum et al. 2004; Karlson et al. 2007). N. melanostomus is a very important food source for Great Cormorants (Phalacrocorax carbo), constituting at least 60% of the fish eaten by this bird (Bzoma and Stępniewicz


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2001). In the Gulf of Gdansk, cormorants have shifted their diet from eel (Anguilla anguilla) and sprat (Sprattus sprattus) to N. melanostomus, resulting in population increases in eel and sprat. The planktivorous sprat has reduced the abundance of zooplankton and hence its grazing pressure on phytoplankton, which has resulted in an increased biomass of the latter (Corkum et al. 2004). N. melanostomus is a euryhaline species and from the colonised brackish waters it succesfully invades freshwater habitats where it may constitute a threat to local species (Grabowska et al. 2010; Poos et al. 2010).

Oncorhynchus mykiss: The rainbow trout O. mykiss exists as a diadromous (often called steelhead) and landlocked freshwater species, although marine populations exist in some European coastal areas (Jonsson 2011). Although the species can have substantial negative impacts (hybridization, disease transmission, predation and competition with native species) in freshwater bodies, no documented negative impacts have been reported in the European marine environment (Jonsson 2011).

Oncorhynchus tshawytscha: The Chinook salmon is not invasive in Europe. According to Crawford and Muir (2008) attempts to introduce this species in European countries were never successful and were soon abandoned. It seems that the species has no impact on ecosystem services and biodiversity in European Seas.

Plotosus lineatus: The striped eel catfish is a venomous fish that arrived in the Mediterranean in 2001 (Golani 2002) and rapidly established a large population (Edelist et al. 2012). Dozens of injuries to fishermen and beachgoers by this species have been reported in Israel with temporary paralysis of limbs and hospitalizations (Gweta et al. 2008; Golani 2010). P. lineatus is taken as by-catch and is discarded. It has a significant commercial value in the aquarium industry (Otero et al. 2013), however no such market seems to have been developed in the Mediterranean.

Salvelinus fontinalis: The brook trout is mainly a freshwater species but diadromous populations occur where some fish may remain at sea for up to three months, especially during spring as stream temperatures rise (Smith and Saunders 1958). In the freshwater environment the species has a high impact (negative: it competes and preys on native fish such as other salmonids for food and cover; it replaces native brown trout Salmo trutta; it puts high pressure on many invertebrate populations through predation; it alters nutrient cycles; positive: important for recreational fishing) (DAISIE 2013) but no impact on biodiversity or ecosystem services has been reported for the marine environment of Europe.

Saurida undosquamis: Since the mid-1950s the brushtooth lizardfish has become commercially important, constituting a substantial portion of the trawl catches in many areas of the eastern Mediterranean (Galil 2007a and references therein). This sudden increase came at the expense of the native hake, Merluccius merluccius (Linnaeus, 1758), and the native lizardfish Synodus saurus, which were reported to be displaced (Ben Yami and Glaser 1974; Otero et al. 2013).

Siganus luridus & Siganus rivulatus: The dusky spinefoot and marbled spinefoot are considered to be high-impact invasive species in the eastern Mediterranean Sea (Katsanevakis et al. 2009; Katsanevakis 2011). They have become dominant in many coastal areas (e.g. Bariche et al. 2004; Katsanevakis 2011; Sala et al. 2011E; Thessalou-Legaki et al. 2012), outcompeting the main native herbivores, Sparisoma cretense (Linnaeus, 1758) and Sarpa salpa (Linnaeus, 1758) (Bariche et al. 2004; Kalogirou et al. 2012), and altering the community structure and the native food web of the rocky infralittoral zone (Sala et al. 2011E; Giakoumi 2014). Based on a caging experiment, Sala et al. (2011)E concluded that S. luridus and S. rivulatus were able to create and maintain barrens (rocky areas almost devoid of erect algae) and contribute to the transformation of the ecosystem from one dominated by lush and diverse brown algal forests to another dominated by bare rock. Some of these algal forests, such as Cystoseira spp. forests, are ecologically very important as nurseries for a number of littoral fish species. These Cystoseira forests are currently considered to be a threatened habitat in several regions of the Mediterranean (Otero et al. 2013). Hence, ecosystem services provided by many sublittoral biotopes, especially communities of sublittoral algae on rocky bottoms, i.e. food, biotic materials, climate regulation, water purification, cognitive benefits, recreation, symbolic and aesthetic values, and life cycle maintenance (Salomidi et al. 2012), are impacted. The species are edible and are caught by trammel nets and gillnets; they are marketed in many Mediterranean countries. In 2008, S. luridus and S. rivulatus represented 4.6% in weight of the total catch of the artisanal fisheries in Cyprus (Katsanevakis et al. 2009). S. rivulatus was cultured on an experimental scale in Egypt, Israel and Cyprus (DAISIE 2013).


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