Ocean acidification, more than warming or heatwaves, constrains shoaling behaviour in a range-extending fish through...

Social context is a critical yet underexplored determinant of behavioural resilience to climate change. Group living can buffer individuals against environmental stress through enhanced vigilance, reduced predation risk and improved foraging efficiency. However, whether these behavioural expressions persist under chronic (warming, acidification) and acute (marine heatwaves) climate stressors remains unclear. Using natural climate analogues spanning present-day, ocean warming and combined warming-acidification reefs, we quantified how shoal size influences behavioural expression in a range-extending reef fish (Pomacentrus coelestis). Across all climate conditions, fish in larger shoals consistently exhibited higher foraging and activity levels and reduced risk-avoidance behaviours, whereas direct effects of warming, acidification and heatwaves on behaviour were negligible. In contrast, ocean acidification most likely constrained collective behaviour indirectly by simplifying benthic habitats, where fish densities were 84% lower than at the warming reef, resulting in shoals that were up to 79% smaller than the Warming and Control reefs. Combined, our data suggest that shoal size mediates behavioural expression between foraging and predator avoidance and that acidification-driven habitat simplification can alter behavioural expression indirectly by reducing fish densities and the formation of large shoals. We conclude that climate change can indirectly modify behavioural expression in shoal-forming fishes through habitat-driven erosion of social structure.