Natural and anthropogenic factors drive large-scale freshwater fish invasions.

We analyzed the large-scale drivers of biological invasions using freshwater fish in a Mediterranean country as a test case, and considering the contribution of single species to the overall invasion pattern. Using Boosted Regression Tree (BRT) models, variation partitioning and Redundancy Analysis (RDA), we found that human factors (especially eutrophication) and climate (especially temperature) were significant drivers of overall invasion. Geography was also relevant in BRT and RDA analysis, both at the overall invasion and the single species level. Only variation partitioning suggested that land use was the second most significant driver group, with considerable overlap between different invasion drivers and only land use and human factors standing out for single effects. There was general accordance both between different analyses, and between invasion outcomes at the overall and the species level, as most invasive species share similar ecological traits and prefer lowland river stretches. Human-mediated eutrophication was the most relevant invasion driver, but the role of geography and climate was at least equally important in explaining freshwater fish invasions. Overall, human factors were less prominent than natural factors in driving the spread and prevalence of invasion, and the species spearheading it.