Bigger Is Better: Characteristics of Round Gobies Forming an Invasion Front in the Danube River

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Abstract
Few studies have systematically investigated differences in performance, morphology and parasitic load of invaders at different stages of an invasion. This study analyzed phenotype-environment correlations in a fish invasion from initial absence until establishment in the headwater reach of the second largest European river, the Danube. Here, the round goby (Neogobius melanostomus) formed 73% of the fish abundance and 58% of the fish biomass in rip-rap bank habitats after establishment. The time from invasion until establishment was only about two years, indicating rapid expansion. Founder populations from the invasion front were different from longer established round goby populations in demography, morphology, feeding behaviour, sex ratio and parasitic load, indicating that plasticity in these traits determines invasion success. Competitive ability was mostly dependent on growth/size-related traits rather than on fecundity. As revealed by stable isotope analyses, specimens at the invasion front had a higher trophic position in the food web and seem to benefit from lower food competition. Somatic performance seems to be more important than investment in reproduction during the early stages of the invasion process and upstream-directed range expansion is not caused by out-migrating weak or juvenile individuals that were forced to leave high density areas due to high competition. This mechanism might be true for downstream introductions via drift. Greater abundance and densities of acanthocephalan endoparasites were observed at the invasion front, which contradicts the expectation that invasion success is determined by lower parasitic pressure in newly invaded areas. Overall, the pronounced changes in fish and invertebrate communities with a dominance of alien species suggest invasional meltdown and a shift of the upper Danube River towards a novel ecosystem with species that have greater resistance to goby predation. This seems to contribute to overcoming biological resistance and improve rapidity of dispersal.

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