Movement responses to lethal risk: an integrative analysis of proactive and reactive antipredator behaviours in a large herbivore

Prey species can display antipredation movement behaviours to reduce predation risk including proactive responses to chronic or predictable risk, and reactive responses to acute or unpredictable risk. Thus, at any given time, prey movement choice may reflect the trade-off between proaction and reaction. In previous studies, proaction and reaction have generally been considered separately, which neglects their simultaneous influence on animal decisions. In this study, we analysed how proaction and reaction interact to shape the movements of GPS-collared red deer (Cervus elaphus), in response to human hunting of conspecifics. Our results show that red deer proactively selected canopy cover where and when risk was predictably high. However, when they were unable to avoid risk, canopy cover was no longer selected, but only modulated a reactive response along a freeze-to-escape continuum. This reaction was even more evident when the environment was unfamiliar, underlining the importance of memory in such reaction patterns. Therefore, to our knowledge, for the first time, we describe how proaction and reaction fuse in an antipredation sequence of interconnected movement decisions in a large herbivore, and we lay the foundations for further investigations into the evolutionary origins of similarities and differences between proactive and reactive behaviours.