The frequency, intensity and duration of aquatic heatwaves are increasing due to climate change globally. However, the ecological consequences of heatwaves are still mainly assessed by acute mass mortality events. This emphasis overlooks the less visible but more pervasive fitness costs that persist after acute exposure. Here, we review the cryptic costs of heatwaves and show how sublethal, delayed and transgenerational effects can possibly depress population fitness and destabilize ecosystems even if the immediate mortality observed is limited. We synthesize evidence that aquatic heatwaves impose cryptic costs through four major pathways. First, heatwaves impair physiology by driving overload of bioenergy, oxidative damage, immune dysfunction and imbalance of host-microbiome interactions. These reactions force organisms to prioritize short-term survival and sacrifice self-maintenance and repair. Second, these physiological costs could alter the trajectories of growth, reduction of reproductive investment and trade-offs in stage-specific life-history. Third, behavioral responses caused by thermal stress often create new ecological liabilities by impairing cognition and locomotion, compressing usable habitat and increasing exposure risks to predators, pollutants and human activities. Fourth, parental heatwave exposure can modify offspring performance through altering provision behavior, disrupting microbiomes and causing epigenetic effects. Thus, these adverse effects extend across generations. In this review, we point out the priorities for future research should particularly focus on integrative frameworks that link multi-stressor environments with multigenerational processes. Therefore, assessments of aquatic heatwave impacts should extend beyond immediate resistance during exposure to include post-event recovery, recruitment and future population dynamics. Instead, aquatic heatwaves should be understood as disturbances that can leave lasting demographic and ecosystem legacies even in the absence of obvious die-offs. Therefore, integrating cryptic fitness costs into demographic models, multi-stressor experiments and ecosystem forecasting will be essential for predicting the effect of heatwaves in reshaping aquatic biodiversity in a warming world.




