Quantifying the behavioural consequences of shark ecotourism.

Shark populations globally are facing catastrophic declines. Ecotourism has been posited as a potential solution to many of the issues facing shark conservation, yet increasingly studies suggest that such activity may negatively influence aspects of shark ecology and so further pressure declining populations. Here we combine UAV videography with deep learning algorithms, multivariate statistics and hidden Markov models (HMM) to quantitatively investigate the behavioural consequences of ecotourism in the whale shark (Rhincodon typus). We find that ecotourism increases the probability of sharks being in a disturbed behavioural state, likely increasing energetic expenditure and potentially leading to downstream ecological effects. These results are only recovered when fitting models that account for individual variation in behavioural responses and past behavioural history. Our results demonstrate that behavioural responses to ecotourism are context dependent, as the initial behavioural state is important in determining responses to human activity. We argue that models incorporating individuality and context-dependence should, wherever possible, be incorporated into future studies investigating the ecological impacts of shark ecotourism, which are only likely to increase in importance given the expansion of the industry and the dire conservation status of many shark species.