Intermittent Search, Not Strict Lévy Flight, Evolves under Relaxed Foraging Distribution Constraints.

AbstractThe survival of an animal depends on its success as a forager, and understanding the adaptations that result in successful foraging strategies is an enduring endeavour of behavioral ecology. Random walks are one of the primary mathematical descriptions of foraging behavior. Power law distributions are often used to model random walks, as they can characterize a wide range of behaviors, including Lévy walks. Empirical evidence indicates the prevalence and efficiency of Lévy walks as a foraging strategy, and theoretical work suggests an evolutionary origin. However, previous evolutionary models have assumed a priori that move lengths are drawn from a power law or other families of distributions. Here, we remove this restriction with a model that allows for the evolution of any distribution. Instead of Lévy walks, our model unfailingly results in the evolution of intermittent search, a random walk composed of two disjoint modes-frequent localized walks and infrequent extensive moves-that consistently outcompeted Lévy walks. We also demonstrate that foraging using intermittent search may resemble a Lévy walk because of interactions with the resources within an environment. These extrinsically generated Lévy-like walks belie an underlying behavior and may explain the prevalence of Lévy walks reported in the literature.

The evolutionary maintenance of Lévy flight foraging.

Lévy flight is a type of random walk that characterizes the behaviour of many natural phenomena studied across a multiplicity of academic disciplines; within biology specifically, the behaviour of fish, birds, insects, mollusks, bacteria, plants, slime molds, t-cells, and human populations. The Lévy flight foraging hypothesis states that because Lévy flights can maximize an organism's search efficiency, natural selection should result in Lévy-like behaviour. Empirical and theoretical research has provided ample evidence of Lévy walks in both extinct and extant species, and its efficiency across models with a diversity of resource distributions. However, no model has addressed the maintenance of Lévy flight foraging through evolutionary processes, and existing models lack ecological breadth. We use numerical simulations, including lineage-based models of evolution with a distribution of move lengths as a variable and heritable trait, to test the Lévy flight foraging hypothesis. We include biological and ecological contexts such as population size, searching costs, lifespan, resource distribution, speed, and consider both energy accumulated at the end of a lifespan and averaged over a lifespan. We demonstrate that selection often results in Lévy-like behaviour, although conditional; smaller populations, longer searches, and low searching costs increase the fitness of Lévy-like behaviour relative to Brownian behaviour. Interestingly, our results also evidence a bet-hedging strategy; Lévy-like behaviour reduces fitness variance, thus maximizing geometric mean fitness over multiple generations.