Summary 1.Pathways describe the processes that result in the introduction of alien species from one location to another. A framework is proposed to facilitate the comparative analysis of invasion pathways by a wide range of taxa in both terrestrial and aquatic ecosystems. Comparisons with a range of data helped identify existing gaps in current knowledge of pathways and highlight the limitations of existing legislation to manage introductions of alien species. The scheme aims for universality but uses the European Union as a case study for the regulatory perspectives. 2. Alien species may arrive and enter a new region through three broad mechanisms: importation of a commodity, arrival of a transport vector, and/or natural spread from a neighbouring region where the species is itself alien. These three mechanisms result in six principal pathways: release, escape, contaminant, stowaway, corridor and unaided. 3. Alien species transported as commodities may be introduced as a deliberate release or as an escape from captivity. Many species are not intentionally transported but arrive as a contaminant of a commodity, for example pathogens and pests. Stowaways are directly associated with human transport but arrive independently of a specific commodity, for example organisms transported in ballast water, cargo and airfreight. The corridor pathway highlights the role transport infrastructures play in the introduction of alien species. The unaided pathway describes situations where natural spread results in alien species arriving into a new region from a donor region where it is also alien. 4. Vertebrate pathways tend to be characterized as deliberate releases, invertebrates as contaminants and plants as escapes. Pathogenic micro-organisms and fungi are generally introduced as contaminants of their hosts. The corridor and unaided pathways are often ignored in pathway assessments but warrant further detailed consideration. 5. Synthesis and applications. Intentional releases and escapes should be straightforward to monitor and regulate but, in practice, developing legislation has proved difficult. New introductions continue to occur through contaminant, stowaway, corridor and unaided pathways. These pathways represent special challenges for management and legislation. The present framework should enable these trends to be monitored more clearly and hopefully lead to the development of appropriate regulations or codes of practice to stem the number of future introductions.

There are about 100 nonindigenous species recorded in the Baltic Sea. Invasive species have resulted in major changes in nearshore ecosystems, especially in coastal lagoons and inlets that can be identified as "centres of xenodiversity". Fewer than 70 of these species have established reproducing populations. Dominant invasion vectors include unintentional introductions via ballast water, tank sediments, and hull fouling, aquaculture, and the construction of canals that have facilitated active or passive natural dispersal. Of the approximately 60 unintentionally introduced species with a known invasion history, 38 are transoceanic (including 19 Atlantic species of American origin) and 18 of Ponto-Caspian origin. Species that have caused economic damage to fisheries, shipping, and industry include the hydrozoan Cordylophora caspia, the barnacle Balanus improvisus, the cladoceran Cercopagis pengoi, and the bivalve Dreissena polymorpha. The Baltic Sea has served as a secondary source of nonindigenous species to the North American Great Lakes. Further study is warranted to quantify large-scale ecosystem changes in the Baltic associated with establishment and population growth of nonindigenous species and to prevent future invasions.

Taking into account the continuous increase in freshwater introductions, and to support the recent European legislation on invasive alien species, the identification of priority pathways and gateways of introductions is of utmost importance to develop adequate control strategies. The aim of this paper was to analyse the main pathways and gateways of introductions of freshwater alien species in Europe. Based on a thorough review of the scientific and grey literature, information on pathways, country and year of initial introduction of all freshwater alien species in Europe, was retrieved. The spatial and temporal patterns and trends of biological invasions in freshwater ecosystems in Europe, in relation to different pathways, were assessed. Our results pinpoint the major importance of aquaculture, pet/aquarium trade and stocking activities as pathways of introduction of freshwater alien species in Europe. For species native to some European countries, shipping and inland canals were the most important pathways, highly responsible for the entry of many harmful species. Germany, the United Kingdom and Italy were the main entry gateways of freshwater alien species in Europe. We found a geographical pattern related to some pathways of introduction in Europe: introductions through inland canals were concentrated in Central/North-eastern Europe, while introductions through pet/terrarium/aquarium trade were mainly observed in Central/Western Europe. While Chordata species entered Europe mainly through the three major above mentioned pathways, many harmful Arthropoda and Mollusca entered through shipping and inland canals. The information gathered in this study clearly indicates the entry routes that should be prioritised by Member States, for which stronger control and management actions should be implemented and prevention efforts concentrated under the scope of the related new EU Regulation.

Over the past century, the potential for aquatic species to expand their ranges in Europe has been enhanced both as a result of the construction of new canals and because of increased international trade. A complex network of inland waterways now connects some previously isolated catchments in southern (Caspian, Azov, Black, Mediterranean seas) and northern (Baltic, North, Wadden, White seas) Europe, and these waterways act as corridors for nonnative species invasions. We have developed a conceptual risk assessment model for invasive alien species introductions via European inland waterways, with specific protocols that focus on the development of environmental indicators within the socioeconomic context of the driving forces-pressures-state-impact-response framework. The risk assessment protocols and water quality indicators on alien species were tested for selected ecosystems within 3 main European invasion corridors, and these can be recommended for application as part of the Common Implementation Strategy of the European Commission Water Framework Directive, which aims to provide a holistic risk-based management of European river basins. The conceptual structure of the online Risk Assessment Toolkit for aquatic invasive alien species is provided and includes 3 main interlinked components: online risk assessment protocols, an early warning system, and an information transmitter for risk communication to end users.

The distribution and abundance of the invasive ctenophore Mnemiopsis leidyi in the Bornholm Basin, an important spawning ground of several fish stocks, and in adjacent areas in the central Baltic Sea was studied in November 2007. The study showed that M. leidyi were relatively small (body length 18.6 ± 7.6 mm) and they were patchily distributed over a large part of the investigated area. Specimens were found on 68 and 59% of stations sampled with a Bongo net (n=39) and an Isaac-Kidd midwater trawl (n=51), respectively. Vertically, the highest densities of M. leidyi occurred at 40 to 60 m around the halocline. Horizontally, the highest abundances were found north and west of Bornholm, but relatively high densities were also observed in the Slupsk Furrow. The mean abundance was 1.58 ± 2.12 ind. m -2 , the peak abundance was 8.92 ind. m -2 , and the average and peak population density were 0.03 ± 0.05 and 0.28 ind. m -3 , respectively. The abundances are low compared to densities recently observed in other areas of the Baltic region (e. g. Limfjorden, Åland Sea) and the estimated predation impact on zooplankton by M. leidyi was negligible in November 2007. However, because of the ctenophore's wide distribution in the central Baltic Sea, its ability for rapid population growth, and its potential influence on fish stocks by competing for food and by preying on fish eggs and newly hatched larvae, close monitoring of the future development of M. leidyi in the Baltic Sea is strongly recommended.

No abstract

The invasion of the zebra mussel, Dreissena polymorpha, into North American waters has resulted in profound ecological disturbances and large monetary losses. This study examined the invasion history and patterns of genetic diversity among endemic and invading populations of zebra mussels using DNA sequences from the mitochondrial cytochrome oxidase I (COI) gene. Patterns of haplotype frequency indicate that all invasive populations of zebra mussels from North America and Europe originated from the Ponto-Caspian Sea region. The distribution of haplotypes was consistent with invasive populations arising from the Black Sea drainage, but could not exclude the possibility of an origin from the Caspian Sea drainage. Similar haplotype frequencies among North American populations of D. polymorpha suggest colonization by a single founding population. There was no evidence of invasive populations arising from tectonic lakes in Turkey, while lakes in Greece and Macedonia contained only Dreissena stankovici. Populations in Turkey might be members of a sibling species complex of D. polymorpha. Ponto-Caspian derived populations of D. polymorpha (theta = 0.0011) and Dreissena bugensis (one haplotype) exhibited low levels of genetic diversity at the COI gene, perhaps as a result of repeated population bottlenecks. In contrast, geographically isolated tectonic lake populations exhibited relatively high levels of genetic diversity (theta = 0.0032 to 0.0134). It is possible that the fluctuating environment of the Ponto-Caspian basin facilitated the colonizing habit of invasive populations of D. polymorpha and D. bugensis. Our findings were concordant with the general trend of destructive freshwater invaders in the Great Lakes arising from the Ponto-Caspian Sea basin.

No abstract