Scientific name parva Common name Stone moroko Broad group Vertebrate Number of and countries wherein the 16: AT, BE, BG, CZ, DE, DK, GR, ES, FR, HU, IT, NL, PL, RO, SK, UK species is currently established Risk Assessment GB NNRA Method https://secure.fera.defra.gov.uk/nonnativespecies/downloadDocument.cfm? Links id=243 Additional EU countries where the species is found (3): Sweden (CABI), Croatia (GISIN), Slovenia (Gozlan et al., 2010).

The species has been recorded in the following: EU member: Austria, Belgium, Bulgaria, Croatia, Czech Republic, Denmark, France, Germany, Greece, Hungary, Italy, Lithuania, Netherlands, Poland, Romania, Slovakia, Spain, Sweden & United Kingdom (19)

It is currently established populations in the following EU member states: 1. Description Austria, Belgium, Bulgaria, Czech Republic, Denmark, France, Germany, (, invasion Greece, Hungary, Italy, Lithuania, Netherlands, Poland, Romania, Slovakia, history, distribution Spain, Sweden & United Kingdom (18) range (native and introduced), This species has shown signs of invasiveness in the following EU member geographic scope, states: socio-economic Austria, Belgium, Bulgaria, Czech Republic, Denmark, France, Germany, benefits) Greece, Hungary, Italy, Lithuania, Netherlands, Poland, Romania, Slovakia, Spain, Sweden & United Kingdom (18)

It could be established in the following EU Biogeographic area: Atlantic, Alpine, Continental, Pannonian, Mediterranean & Steppic

The species could establish in the future [given current climate] (including those where it is already established) in the following EU MS: Austria, Belgium, Bulgaria, Croatia, Cyprus, Czech Republic, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, Ireland, Italy, Latvia, Lithuania, Luxembourg, Malta, Netherlands, Poland, Portugal, Romania, Slovakia, Slovenia, Spain, Sweden & United Kingdom (28)

It could become invasive in the future [given current climate] (where it is not already established) in: Croatia, Cyprus, Estonia, Finland, Ireland, Latvia, Luxembourg, Malta, Portugal, Slovenia (10)

Socio-economic benefits: Some commercially valuable fish species, such as the Chinese perch, are reported to pray on the topmouth gudgeon in the native range (China, Japan) (FishBase, http://www.fishbase.org).

Feed item for commercially important species in aquaculture. Considered a model species for research. The primary introduction pathways, in order of importance, are aquaculture, mainly associated to the species’ hitch-hiking with Chinese 4. Has the capacity to carp species and common carp Cyprinus carpio (65%), recreational fishing assess multiple (22%), ornamental fish trade (9%) and natural dispersal (1%). However, pathways of entry and natural dispersal represents the main secondary pathway (72%) followed spread in the by angling (25%) and the ornamental fish trade (3%). Based on the assessment, both number of fish movements and introductions during the 1970 and early intentional and 1980’s across European countries, and in particular of Chinese carp unintentional species, it is likely that the actual distribution of P. parva in its invasive range reflects a combination of ‘stepping–stone’ and ‘diffusion’ dispersal models (Gozlan et al., 2010). Provisioning: Food (aquaculture) Inter-specific competition for food between P. parva and other cyprinid fish species resulted in food web structure changes causing increased economic costs for carp aquaculture ponds in the Czech Republic (Gozlan 6. Can broadly assess et al., 2010) . environmental impact with respect to Regulating: Pest and disease regulation ecosystem services Transmission of parasites – the species has an important role in the spread of diseases and parasites as a healthy carrier for a number of pathogens (Gozlan et al., 2010).

The only reported parasite specific to P. parva that has been reported is Dactylogyrus squameus and this has facilitated their dispersal to regions in Kazakhstan, Tajikistan, Uzbekistan, the Czech and Slovak Republics and Italy.

The two most pathogenic parasites associated with P. parva in its invasive range are Anguillicola crassus and the rosette agent (Gozlan et al., 2010, Gozlan et al., 2005). Anguillicola crassus is a parasitic nematode that occupies the swimbladder of eels with the capacity to cause high eel mortalities; P. parva acts as an intermediate host. In a location in France, 35% of P. parva were infected with A. crassus (Cesco et al., 2001). The identification of P. parva as a healthy carrier for the intracellular parasite S. destruens is a concern as this pathogen has been responsible for mass mortalities of salmonid fishes in the USA (Arkush et al., 2003) and has since been associated with the decline of native European fish species including sunbleak (Heckel 1843). Although the origin of S. destruens in Europe remains unclear (Gozlan et al., 2010), it may have arrived with P. parva and consequently, the natural dispersal of P. parva throughout Eurasia may have facilitated their spread, posing a potential thread to cyprinid and salmonid populations.

Supporting: Community, foodweb a) Predation and parasitism (Gozlan et al., 2010) In water bodies of China and Germany, P. parva were reported to feed on eggs and larvae of native fish species. They are also reported to be a facultative parasite on other fish species when kept in high densities. Such behaviour was observed in aquaculture ponds of Moldavia where P. parva >1 year old were causing injuries reaching the musculature in H. molitrix, A. nobilis and C. idella, although feeding on tubificid worms (Tubifex spp.) was also observed. b) Competition for food with native fish species Inter-specific competition for food between P. parva and native fish species has been observed in water bodies of Belgium, Bulgaria (T. Trichkova personal communication), Czech Republic (J. Musil personal communication), Germany, Greece and Poland (Gozlan et al., 2010). A stable isotope analysis revealed significant trophic overlap between P. parva, R. rutilus and C. carpio, a shift associated with significantly depressed somatic growth in R. rutilus (Britton et al., 2010b).

Supporting: Production High grazing pressure exerted by dense P. parva populations can also result in changes in the prevalent environmental conditions through top- down effects characterised by increased development of phytoplankton and accelerated eutrophication (Gozlan et al., 2010).

Supporting: Impact on native aquatic species diversity and density. Competes for food, predates upon juvenile native species, disease vector. Can alter ecosystem function and modify habitat. Threat to native/ endangered species.

Provisioning: Reduction in aquaculture productivity.

Regulating: Water use, disease and pest regulation.

Cultural: Loss of recreational angling opportunities and reduced amenity values. There is an evidence for dietary overlap between P. parva and three endemic species in Lake Mikri Prespa (north-western Greece): Pelasgus prespensis (EN, IUCN), Cobitis meridionalis (VU, IUCN) and Alburnoides ohridanus (VU, IUCN).

It was reported that the population of P. prespensis has declined due to the introduction of alien species (P. parva, Rhodeus amarus) (Kottelat & 8. Includes status Freyhof, 2007). (threatened or protected) of species A recent study on the feeding habits of topmouth gudgeon in the Hirfanli or habitat under Reservoir, Central Anatolia, Turkey, reported a broad food preference threat (including fish eggs) and high feeding intensity of the introduced P. parva. Therefore, it was concluded that P. parva may have a competitive pressure on the native fish fauna in the reservoir - Alburnus cf. escherichi population which was dominant in the 1960s has disappeared (locally extinct) in the Hirfanli Reservoir; studies on food overlap between P. parva and the endemic Aphanius danfordii (CR, IUCN) has been undertaken (F. Güler Ekmekçi, personal communication).

The first evidence of the parasite Sphaerothecum destruens being present in wild populations of topmouth gudgeon, with a prevalence of 67 to 74% was reported in the Netherlands. Sympatric populations of known susceptible fish species were found at the sampled sites, including species that feature in the national Red List (LNV 2004) (bitterling Rhodeus amarus and sunbleak Leucaspius delineatus). No information about infection rates of these native species by S. destruens is available yet, nor about the effects on their population size (Spikmans et al., 2013).

Hybridisation between P. parva and sunbleak L. delineatus (LC, IUCN), a threatened freshwater species that is in decline throughout Europe, has been demonstrated by artificial insemination illustrating gamete compatibility between two genres and a potential for natural hybridisation (Gozlan et al., 2010). Tolerance experience: Recent study evaluated the effect of elevated water temperature on an ecosystem after the invasion by topmouth gudgeon in order to create a possible scenario of climate change and the invasion of topmouth gudgeon (Záhorská et al., 2013). Lake Licheńskie, which is part of the cooling system of the Konin and Pątnów power plants, was used as a model. The building of the cooling system caused an increase in the average temperatures from 5 to 7°C. The modifications of hydrological, thermal and trophic conditions impacted adversely the structure and development of zooplankton and fish communities. The coldwater species typical of the region, such as pikeperch, pike and perch disappeared, or 9. Includes possible occurred at very low densities. On the other hand, submerged vegetation effects of climate formed by the invasive aquatic plants Najas marina and Vallisneria spiralis change in the became abundant and the relative density of topmouth gudgeon foreseeable future increased. The authors assumed that because of the species high phenotypic plasticity in all parameters, from reproduction to morphology, its population would be successful even if the climate changes radically (Záhorská et al., 2013).

Observation: 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 (Gozlan et al., 2010).

Climate matching: increase in climate suitability by 2050 is predicted (Britton et al., 2010a). The predictive use of climate-matching models and an air and water temperature regression model suggested that there are six non-native fishes currently persistent but not established in England and Wales whose establishment and subsequent invasion would benefit substantially from the predicted warming temperatures. These included the common carp Cyprinus carpio and European catfish Silurus glanis, fishes that also exert a relatively high propagule pressure through stocking to support angling and whose spatial distribution is currently increasing significantly, including in open systems. Arkush KD, Mendoza L, Adkison MA, Hedrick RP. 2003. Observations on the life stages of Sphaerothecum destruens ng, n. sp., a mesomycetozoean fish pathogen formally referred to as the rosette agent. Journal of Eukaryotic Microbiology 50: 430-438. Britton J, Cucherousset J, Davies G, Godard M, Copp G. 2010. Non‐ native fishes and climate change: predicting species responses to warming temperatures in a temperate region. Freshwater Biology 55: 1130-1141. Britton JR, Davies GD, Harrod C. 2010. Trophic interactions and consequent impacts of the invasive fish Pseudorasbora parva in a native aquatic foodweb: a field investigation in the UK. Biological Invasions 12: 1533-1542. Cesco H, Lambert A, Crivelli A. 2001. Is Pseudorasbora parva, an invasive fish species (Pisces, ), a new agent of anguillicolosis in France? Parasite 8: 75-76. Gozlan RE, Andreou D, Asaeda T, Beyer K, Bouhadad R, Burnard D, Caiola 11. Documents N, Cakic P, Djikanovic V, Esmaeili HR. 2010. Pan‐continental information sources invasion of Pseudorasbora parva: towards a better understanding of freshwater fish invasions. Fish and Fisheries 11: 315-340. Gozlan RE, St-Hilaire S, Feist SW, Martin P, Kent ML. 2005. Biodiversity: disease threat to European fish. Nature 435: 1046-1046. Kottelat M, Freyhof J. 2007. Handbook of European freshwater fishes. Publications Kottelat Cornol. Spikmans F, van Tongeren T, van Alen TA, van der Velde G, den Camp H. 2013. High prevalence of the parasite Sphaerothecum destruens in the invasive topmouth gudgeon Pseudorasbora parva in the Netherlands, a potential threat to native freshwater fish. Aquat. Invasions 8: 355-360. Záhorská E, Balážová M, Šúrová M. 2013. Morphology, sexual dimorphism and size at maturation in topmouth gudgeon (Pseudorasbora parva) from the heated Lake Licheńskie (Poland). Knowledge and Management of Aquatic Ecosystems: 07.

Teodora Trichkova Main experts Merike Linnamagi Belinda Gallardo Other contributing Olaf Booy experts Guler Ekmekci The topmouth gudgeon Pseudorasbora parva is widely spread in Europe, already reported from 19 EU countries.

Notes In order to fill the information gaps, recent data and data from other regions in Europe on the pathways of introduction, the impact on ecosystem services, impact on protected species and habitats, and results of studies on the effects of climate change are added. Outcome Compliant