A theory for investment across defences triggered at different stages of a predator-prey encounter

We introduce a general theoretical description of a combination of defences acting sequentially at different stages in the predatory sequence in order to make predictions about how animal prey should best allocate investment across different defensive stages. We predict that defensive investment will often be concentrated at stages early in the interaction between a predator individual and the prey (especially if investment is concentrated in only one defence, then it will be in the first defence). Key to making this prediction is the assumption that there is a cost to a prey when it has a defence tested by an enemy, for example because this incurs costs of deployment or tested costs as a defence is exposed to the enemies; and the assumption that the investment functions are the same among defences. But if investment functions are different across defences (e.g. the investment efficiency in making resources into defences is higher in later defences than in earlier defences), then the contrary could happen. The framework we propose can be applied to other victim-exploiter systems, such as insect herbivores feeding on plant tissues. This leads us to propose a novel explanation for the observation that herbivory damage is often not well explained by variation in concentrations of toxic plant secondary metabolites. We compare our general theoretical structure with related examples in the literature, and conclude that coevolutionary approaches will be profitable in future work. (C) 2019 Elsevier Ltd. All rights reserved.