Beyond establishment: incorporating physiological performance into predictions of invasion risk

Biological invasions are a major driver of global change, reshaping ecosystems and threatening biodiversity worldwide. Anticipating where invaders will establish and where they will exert the strongest ecological impacts are key challenges for early detection and targeted management. Although Species Distribution Models (SDMs) are widely used to forecast biological invasions, they often provide uncertain estimates of establishment ranges and limited insight into invader performance, making it difficult to anticipate ecological impacts. Here, we address these limitations by integrating physiological information on invader performance with SDMs to identify regions of high invasion risk. Using the brown alga Rugulopteryx okamurae, one of the most prominent marine invaders in Europe, we first test alternative hypotheses of northern establishment limits: (i) a cold-survival constraint driven by winter temperatures and (ii) a growth constraint derived from the species' thermal performance. To identify the more likely scenario, we combine cold-tolerance experiments with seasonal growth comparisons between the invader and a native macroalga Dictyota dichotoma, whose established distribution allows physiological performance to be directly related to realised presence. Finally, we project seasonal growth of the invader across the predicted establishment range as a proxy for biomass accumulation and potential ecological impacts. Our results indicate that northern limit in Europe will be more likely constrained by winter survival rather than growth, extending the potential establishment range of Rugulopteryx to mid-Norway. In contrast, the highest impacts are likely to remain concentrated in southern Europe, where thermal conditions sustain high year-round growth. Overall, our approach illustrates how understanding the physiological response of invaders to their environment can improve the interpretation of SDM outputs and help identify areas at greatest risk of impact within their potential establishment range.