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Chronic warming, not heatwaves, predicts tropical fish colonization in the Mediterranean

Preprint Created on 08 Sep 2026 bioRxiv

The opening of the Suez Canal abruptly reconnected the Red Sea and the Mediterranean Sea, collapsing a geological barrier that had separated their biotas for millions of years. This exchange offers a natural experiment to determine whether tropical species' spread at their cold-edge range is driven by acute thermal shocks (marine heatwaves) or by progressive easing of physiological constraints under sustained warming. We analyzed 1,059 first-arrival time series for 75 non-indigenous fish species across the Mediterranean, explicitly accounting for progressive salinification, oceanographic connectivity, and historical canal enlargement. Our models demonstrate that colonization dynamics are overwhelmingly dominated by ecological momentum-the time-dependent geographic spread of populations-and facilitated by chronic background warming. Marine heatwaves explain negligible variance and show no detectable effect on colonization success. Instead, warming operates as a stage-dependent continuous climatic facilitator, accelerating post-establishment expansion, rather than triggering introductions. Thus, this shift toward warm-adapted communities is governed by chronic warming, with the Suez Canal triggering the reorganization of Mediterranean biogeography.

Chaikin, S., Vagenas, G., Matias, M., Galanidi, M., Zenetos, A., Araujo, M.

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