Globalization of trade and travel has facilitated the spread of non-native species across the earth. A proportion of these species become established and cause serious environmental, economic, and human health impacts. These species are referred to as invasive, and are now recognized as one of the major drivers of biodiversity change across the globe. As a long-time centre for trade, Europe has seen the introduction and subsequent establishment of at least several thousand non-native species. These range in taxonomy from viruses and bacteria to fungi, plants, and animals. Although invasive species cause major negative impacts across all regions of Europe, they also offer scientists the opportunity to develop and test theory about how species enter and leave communities, how non-native and native species interact with each other, and how different types of species affect ecosystem functions. For these reasons, there has been recent growth in the field of invasion biology as scientists work to understand the process of invasion, the changes that invasive species cause to their recipient ecosystems, and the ways that the problems of invasive species can be reduced. This review covers the process and drivers of species invasions in Europe, the socio-economic factors that make some regions particularly strongly invaded, and the ecological factors that make some species particularly invasive. We describe the impacts of invasive species in Europe, the difficulties involved in reducing these impacts, and explain the policy options currently being considered. We outline the reasons that invasive species create unique policy challenges, and suggest some rules of thumb for designing and implementing management programs. If new management programs are not enacted in Europe, it is inevitable that more invasive species will arrive, and that the total economic, environmental, and human health impacts from these species will continue to grow.

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Invasive species in Europe: ecology, status, and policy

In this paper it is argued, using examples of disease emergence in aquatic animals in Europe, that the introduction of non-native species drives disease emergence by both extending the geographic range of parasites and pathogens and facilitating host-switching. Enteric red mouth disease and infectious haematopoietic necrosis of salmonids have extended their geographic range from North America to Europe with the import of live fish (Pimephales promelas) and rainbow trout eggs, respectively. Host-switching results in disease emergence when previously unidentified commensal organisms or known pathogen switch to new naive hosts. The most serious endemic diseases of wild aquatic animals in Europe in recent years can be traced to the introduction of non-native species. Across Europe dramatic populations declines have occurred in native crayfish (e.g. Astacus astacus), oysters (Ostrea edulis) and eels (Anguilla anguilla), all which can be attributed, in varying degrees, to diseases (crayfish plague, Bonamia ostreae and Anguillicoloides crassus, respectively) introduced with non-native species. The severe adverse effects at a population level can be attributed to the lack of immunity in the new hosts. The impact of parasites more recently introduced to Europe, Sphaerothecum destruens (the rosette agent), and Batrachochytrium dendrobatidis, have yet to be fully determined. Both are generalists, with wide host ranges, and may present serious threats to native species. Aquaculture is the key driver for the introduction of non-native species. Most farming systems allow pathogen exchange between farmed and wild populations which underpins host-switching. Subsequently movements of animals between farms may result in the spread of newly emerged diseases. The introduction of non-native aquatic animals drives disease emergence, thus the ex-ante assessment of these hazards is severely limited. Generic risk mitigation measures (e.g. movement of disinfected eggs in place of live animals) and improved methods for rapid detection of new diseases are vital.

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Non-native aquatic animals introductions have driven disease emergence in Europe

Nonânative fishes are often blamed for exerting adverse ecological impacts on native fishes, but the evidence is circumstantial and the true impacts remain largely unknown. This dilemma is particularly acute for freshwater ecosystems of the Mediterranean Region, where a high level of endemism (a biodiversity hotspot) makes native fish extinctions much more likely because of their small natural ranges. The aim of this paper is to review nonânative fish impacts, from genes to ecosystem level, in the fresh waters of the Mediterranean Region and identify gaps in knowledge that currently hamper effective conservation and management of native species and ecosystems. Most studies reviewed reported potential impacts rather than realised impact mechanisms, and these were conducted predominantly in the European part of the Mediterranean Region. No studies of realised impacts were found for three of the 15 most introduced fishes, which emphasises the need for evidenceâbased studies of nonânative species impacts.

Non‐native fish impacts on Mediterranean freshwater ecosystems: current knowledge and research needs

Using the electrofishing database of the French National Agency for Water and Aquatic Environment (Onema), the time trends of 48 freshwater fish taxa at 590 sites monitored for at least 8 years from 1990 to 2009 were investigated. The results demonstrated that species richness increased steadily from the beginning of the monitoring period. This is congruent with the finding that the number of species displaying a significant increase in spatial distribution or abundance was greater than those showing a significant decrease. Some species, however, had declined both in occurrence and abundance, e.g. tench Tinca tinca, common bream Abramis brama, brown trout Salmo trutta and European eel Anguilla anguilla. The species showing the most spectacular colonization were non-native, e.g. topmouth gudgeon Pseudorasbora parva, wels catfish Silurus glanis and asp Aspius aspius. The time trends in population density were related to the maximal body size, habitat requirement, occurrence and abundance and the status (i.e. native or exotic) but not to the spawning temperature.

Time trends in fish populations in metropolitan France: insights from national monitoring data

Summary 1. Ecological theory attempts to predict how impacts for native species arise from biological invasions. A fundamental question centres on the feeding interactions of invasive and native species: whether invasion will result in increased interspecific competition, which would result in negative consequences for the competing species, or trophic niche divergence, which would facilitate the invader’s integration into the community and their coexistence with native species. 2. Here, the feeding interactions of a highly invasive fish, topmouth gudgeon Pseudorasbora parva, with three native and functionally similar fishes were studied to determine whether patterns of either niche overlap or divergence detected in mesocosm experiments were apparent between the species at larger spatial scales. Using stable isotope analysis, their feeding relationships were assessed initially in the mesocosms (1000 L) and then in small ponds ( 600 m 2 ). 3. In the mesocosms, a consistent pattern of trophic niche divergence was evident between the sympatric fishes, with niches shifting further apart in isotopic space than suggested in allopatry, revealing that sharing of food resources was limited. Sympatric P. parva also had a smaller niche than their allopatric populations. 4. In eight small ponds where P. parva had coexisted for several years with at least one of the fish species used in the mesocosms, strong patterns of niche differentiation were also apparent, with P. parva always at a lower trophic position than the other fishes, as also occurred in the mesocosms. Where these fishes were sympatric within more complex fish communities in the large ponds, similar patterns were also apparent, with strong evidence of trophic niche differentiation. 5. Aspects of the ecological impacts of P. parva invasion for native communities in larger ponds were consistent with those in the mesocosm experiments. Their invasion resulted in divergence in trophic niches, partly due to their reduced niche widths when in sympatry with other species, facilitating their coexistence in invaded ecosystems. Our study highlights the utility of controlled mesocosm studies for predicting the trophic relationships that can develop from introductions of non-native species into more complex ecosystems and at larger spatial scales.

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Patterns of trophic niche divergence between invasive and native fishes in wild communities are predictable from mesocosm studies

In recent years, policy-makers have sought the development of appropriate tools to prevent and manage introductions of invasive species. However, these tools are not well suited for introductions of non-target species that are unknowingly released alongside intentionally-introduced species. The most compelling example of such invasion is arguably the topmouth gudgeon Pseudorasbora parva, a small cyprinid species originating from East Asia. A combination of sociological, economical and biological factors has fuelled their rapid invasion since the 1960s; 32 countries (from Central Asia to North Africa) have been invaded in less than 50 years. Based on a combination of monitoring surveys (2535 populations sampled) and literature reviews, this paper aims to quantify and characterise important invasion parameters, such as pathways of introduction, time between introduction and detection, lag phase and plasticity of life history traits. Every decade, five new countries have reported P. parva introduction, mainly resulting from the movement of Chinese carps for fish farming. The mean detection period after first introduction was 4 years, a duration insufficient to prevent their pan-continental invasion. High phenotypic plasticity in fitness related traits such as growth, early maturity, fecundity, reproductive behaviour and the ability to cope with novel pathogens has predisposed P. parva to being a strong invader. The Pseudorasbora parva invasion has provided quantitative data for the development of 1) early warning systems across different spatial scales; 2) rapid eradication programmes prior to natural spread in open systems and 3) sound risk assessments with emphasis on plasticity of life history traits.

Chun Guang Zhang

Papers

Pan-continental invasion of Pseudorasbora parva: towards a better understanding of freshwater fish invasions