A good understanding of the mechanisms and magnitude of impact of invasive alien species on ecosystem services and biodiversity is a prerequisite for efficient prioritization of actions for prevention of new invasions or for mitigation measures. In this review, we have identified marine alien species of high-impact on ecosystem services and biodiversity in the European seas, classified the mechanisms of impact, commented on the methods applied for assessing impact and the related inferential strength, and reported gaps in available information. Furthermore, we have proposed an updated inventory of 86 marine species in Europe, representing 12 phyla, with documented high impact on ecosystem services or biodiversity. Food provision was the ecosystem service that was impacted by the highest number of alien species (in terms of both positive and negative impacts). Following food provision, the ecosystem services that were negatively affected by the highest number of species were ocean nourishment, recreation and tourism, and water purification, while the ecosystem services that were more often positively impacted were water purification, cognitive benefits, and climate regulation. In many cases, marine aliens impact keystone/protected species and habitats. Almost 30% of the assessed species affected entire ecosystem processes or wider ecosystem function, more often in a negative way. Fifty of the assessed species have been reported to be ecosystem engineers, fundamentally modifying, creating or defining habitats by altering their physical or chemical properties. Positive impacts of alien species are probably underestimated, as there is a persistent perception bias against alien species. We conclude that the “native good, alien bad” philosophy is a misconception and the role of most of the alien species in marine ecosystems is rather complex. Among the species herein assessed as high-impact species, 15 had only negative and 8 only positive impacts; for the majority (64 species) both negative and positive impacts have been reported, and the overall balance is often unknown. Although there is no doubt that invasive species have modified marine ecosystems, evidence for most of the reported impacts is weak, as it is based on expert judgement or dubious correlations, while only ~10% of the reported impacts were inferred via manipulative or natural experiments. A need for stronger inference is evident, to improve our knowledge base of marine biological invasions and better inform managers.JRC.H.1-Water Resource

More than 60 marine non-indigenous species (NIS) have been removed from previous lists and 84 species have been added, bringing the total to 986 alien species in the Mediterranean [775 in the eastern Mediterranean (EMED), 249 in the central Mediterranean (CMED), 190 in the Adriatic Sea (ADRIA) and 308 in the western Mediterranean (WMED)]. There were 48 new entries since 2011 which can be interpreted as approximately one new entry every two weeks. The number of alien species continues to increase, by 2-3 species per year for macrophytes, molluscs and polychaetes, 3-4 species per year for crustaceans, and 6 species per year for fish. The dominant group among alien species is molluscs (with 215 species), followed by crustaceans (159) and polychaetes (132). Macrophytes are the leading group of NIS in the ADRIA and the WMED, reaching 26-30% of all aliens, whereas in the EMED they barely constitute 10% of the introductions. In the EMED, molluscs are the most species-rich group, followed by crustaceans, fish and polychaetes. More than half (54%) of the marine alien species in the Mediterranean were probably introduced by corridors (mainly Suez). Shipping is blamed directly for the introduction of only 12 species, whereas it is assumed to be the most likely pathway of introduction (via ballasts or fouling) of another 300 species. For approximately 100 species shipping is a probable pathway along with the Suez Canal and/or aquaculture. Approximately 20 species have been introduced with certainty via aquaculture, while >50 species (mostly macroalgae), occurring in the vicinity of oyster farms, are assumed to be introduced accidentally as contaminants of imported species. A total of 18 species are assumed to have been introduced by the aquarium trade. Lessepsian species decline westwards, while the reverse pattern is evident for ship-mediated species and for those introduced with aquaculture. There is an increasing trend in new introductions via the Suez Canal and via shipping.

The state-of-art on alien species in the Mediterranean Sea is presented, making distinctions among the four subregions defined in the EU Marine Strategy Framework Directive: (i) the Western Mediterranean Sea (WMED); (ii) the Central Mediterranean Sea (CMED); (iii) the Adriatic Sea (ADRIA); and (iv) the Eastern Mediterranean Sea (EMED). The updated checklist (December 2010) of marine alien species within each subregion, along with their acclimatization status and origin, is provided. A total of 955 alien species is known in the Mediterranean, the vast majority of them having being introduced in the EMED (718), less in the WMED (328) and CMED (267) and least in the Adriatic (171). Of these, 535 species (56%) are established in at least one area.Despite the collective effort of experts who attempted in this work, the number of introduced species remains probably underestimated. Excluding microalgae, for which knowledge is still insufficient, aliens have increased the total species richness of the Mediterranean Sea by 5.9%. This figure should not be directly read as an indication of higher biodiversity, as spreading of so many aliens within the basin is possibly causing biotic homogenization. Thermophilic species, i.e. Indo-Pacific, Indian Ocean, Red Sea, Tropical Atlantic, Tropical Pacific, and circum(sub)tropical, account for 88.4% of the introduced species in the EMED, 72.8% in the CMED, 59.3% in the WMED and 56.1% in the Adriatic. Cold water species, i.e. circumboreal, N Atlantic, and N Pacific, make up a small percentage of the introduced species, ranging between 4.2% and 21.6% and being more numerous in the Adriatic and less so in the EMED.Species that are classified as invasive or potentially invasive are 134 in the whole of the Mediterranean: 108 are present in the EMED, 76 in the CMED, 53 in the Adriatic and 64 in the WMED. The WMED hosts most invasive macrophytes, whereas the EMED has the lion’s share in polychaetes, crustaceans, molluscs and fish.

This collaborative effort by many specialists across the Mediterranean presents an updated annotated list of alien marine species in the Mediterranean Sea. Alien species have been grouped into six broad categories namely established, casual, questionable, cryptogenic, excluded and invasive, and presented in lists of major ecofunctional/taxonomic groups. The establishment success within each group is provided while the questionable and excluded records are commented in brief. A total of 963 alien species have been reported from the Mediterranean until December 2005, 218 of which have been classified as excluded (23%) leaving 745 of the recorded species as valid aliens. Of these 385 (52%) are already well established, 262 (35%) are casual records, while 98 species (13%) remain “questionable” records. The species cited in this work belong mostly to zoobenthos and in particular to Mollusca and Crustacea, while Fish and Phytobenthos are the next two groups which prevail among alien biota in the Mediterranean. The available information depends greatly on the taxonomic group examined. Thus, besides the three groups explicitly addressed in the CIESM atlas series (Fish, Decapoda/Crustacea and Mollusca), which are however updated in the present work, Polychaeta, Phytobenthos, Phytoplankton and Zooplankton are also addressed in this study. Among other zoobenthic taxa sufficiently covered in this study are Echinodermata, Sipuncula, Bryozoa and Ascidiacea. On the contrary, taxa such as Foraminifera, Amphipoda and Isopoda, that are not well studied in the Mediterranean, are insufficiently covered. A gap of knowledge is also noticed in Parasites, which, although ubiquitous and pervasive in marine systems, have been relatively unexplored as to their role in marine invasions. Conclusively the lack of funding purely systematic studies in the region has led to underestimation of the number of aliens in the Mediterranean. Emphasis is put on those species that are current or potential threats to the marine ecosystems, namely the Worst Invasive Alien Species providing their record across major groups.

Spatial priorities for the conservation of three key Mediterranean habitats, i.e. seagrass Posidonia oceanica meadows, coralligenous formations, and marine caves, were determined through a systematic planning approach. Available information on the distribution of these habitats across the entire Mediterranean Sea was compiled to produce basin-scale distribution maps. Conservation targets for each habitat type were set according to European Union guidelines. Surrogates were used to estimate the spatial variation of opportunity cost for commercial, non-commercial fishing, and aquaculture. Marxan conservation planning software was used to evaluate the comparative utility of two planning scenarios: (a) a whole-basin scenario, referring to selection of priority areas across the whole Mediterranean Sea, and (b) an ecoregional scenario, in which priority areas were selected within eight predefined ecoregions. Although both scenarios required approximately the same total area to be protected in order to achieve conservation targets, the opportunity cost differed between them. The whole-basin scenario yielded a lower opportunity cost, but the Alboran Sea ecoregion was not represented and priority areas were predominantly located in the Ionian, Aegean, and Adriatic Seas. In comparison, the ecoregional scenario resulted in a higher representation of ecoregions and a more even distribution of priority areas, albeit with a higher opportunity cost. We suggest that planning at the ecoregional level ensures better representativeness of the selected conservation features and adequate protection of species, functional, and genetic diversity across the basin. While there are several initiatives that identify priority areas in the Mediterranean Sea, our approach is novel as it combines three issues: (a) it is based on the distribution of habitats and not species, which was rarely the case in previous efforts, (b) it considers spatial variability of cost throughout this socioeconomically heterogeneous basin, and (c) it adopts ecoregions as the most appropriate level for large-scale planning.