Assessing Levy walks as models of animal foraging

The hypothesis that the optimal search strategy is a Levy walk (LW) or Levy flight, originally suggested in 1995, has generated an explosion of interest and controversy. Long-standing empirical evidence supporting the LW hypothesis has been overturned, while new models and data are constantly being published. Statistical methods have been criticized and new methods put forward. In parallel with the empirical studies, theoretical search models have been developed. Some theories have been disproved while others remain. Here, we gather together the current state of the art on the role of LWs in optimal foraging theory. We examine the body of theory underpinning the subject. Then we present new results showing that deviations from the idealized one-dimensional search model greatly reduce or remove the advantage of LWs. The search strategy of an LW with exponent mu = 2 is therefore not as robust as is widely thought. We also review the available techniques, and their potential pit-falls, for analysing field data. It is becoming increasingly recognized that there is a wide range of mechanisms that can lead to the apparent observation of power-law patterns. The consequence of this is that the detection of such patterns in field data implies neither that the foragers in question are performing an LW, nor that they have evolved to do so. We conclude that LWs are neither a universal optimal search strategy, nor are they as widespread in nature as was once thought.