In situ effects of human disturbances on coral reef-fish assemblage structure: Temporary and persisting changes are reflected as a result of intensive tourism

Non-lethal human disturbances are often drivers of change in animal population and community structure. To gauge their severity, short-term behaviour (e.g. avoidance and habituation) has been argued to be a sensitive measure. However, many of these behavioural changes may occur only if disturbance-free habitat is readily accessible. In coral-reef fish, we tested whether human disturbances from intensive (i.e. loud music, swimming, snorkelling, splashing and fish feeding by numerous visitors) tourist visitations resulted in assemblage structure shifts led by short-term behaviour. We monitored fish assemblage before, during and after tourist visitations to monitor changes associated with behaviour. Additionally, we monitored two adjacent reefs not visited by tourists because of difficult approach by boat. We posited that if short-term benefits of relocating to disturbance-free habitat outweigh the costs of tolerating disturbances, fish assemblage structure should shift along with tourist visitation levels. By contrast, if sensitive species are unable or unwilling to relocate, we predicted greater levels of assemblage heterogeneity between the visited and control reefs. Our results showed that in situ human visitations led to significant shifts in assemblage structure, resulting from short-term behavioural changes. Additionally, we showed significant between-reefs differences, whereby control reefs were characterised by higher species richness, larger fish sizes and variations in relative trophic guild prevalence. Our results suggest that short-term relocations to adjacent disturbance-free reefs may not mitigate the effects of human disturbances.