The Adaptiveness of Host Behavioural Manipulation Assessed Using Tinbergen's Four Questions

Host organisms show altered phenotypic reactions when parasitised, some of which result from adaptive host manipulation, a phenomenon that has long been debated. Here, we provide an overview and discuss the rationale in distinguishing adaptive versus nonadaptive host behavioural manipulation. We discuss Poulin's criteria of adaptive host behavioural manipulation within the context of Tinbergen's four questions of ethology, while highlighting the importance of both the proximate and evolutionary explanations of such traits. We also provide guidelines for future studies exploring the adaptiveness of host behavioural manipulation. Through this article, we seek to encourage researchers to consider both the proximate and ultimate causes of host behavioural manipulation to infer on the adaptiveness of such traits.