A guide to area-restricted search: a foundational foraging behaviour

Area-restricted search is the capacity to change search effort adaptively in response to resource encounters or expectations, from directional exploration (global, extensive search) to focused exploitation (local, intensive search). This search pattern is used by numerous organisms, from worms and insects to humans, to find various targets, such as food, mates, nests, and other resources. Area-restricted search has been studied for at least 80 years by ecologists, and more recently in the neurological and psychological literature. In general, the conditions promoting this search pattern are: (1) clustered resources; (2) active search (e.g. not a sit-and-wait predator); (3) searcher memory for recent target encounters or expectations; and (4) searcher ignorance about the exact location of targets. Because area-restricted search adapts to resource encounters, the search can be performed at multiple spatial scales. Models and experiments have demonstrated that area-restricted search is superior to alternative search patterns that do not involve a memory of the exact location of the target, such as correlated random walks or Lévy walks/flights. Area-restricted search is triggered by sensory cues whereas concentrated search in the absence of sensory cues is associated with other forms of foraging. Some neural underpinnings of area-restricted search are probably shared across metazoans, suggesting a shared ancestry and a shared solution to a common ecological problem of finding clustered resources. Area-restricted search is also apparent in other domains, such as memory and visual search in humans, which may indicate an exaptation from spatial search to other forms of search. Here, we review these various aspects of area-restricted search, as well as how to identify it, and point to open questions.