Optimal Foraging: A Selective Review of Theory and Tests

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Abstract
Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last 10 yr, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar, in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some currency (Schoener, 1971), usually energy, and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities the models have become known as optimal foraging models; and the theory that embodies them, optimal foraging theory. The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following 4 categories: choice by an animal of which food types to eat, (i.e., optimal diet); choice of which patch type to feed in (i.e., optimal patch choice); optimal allocation of time to different patches; and optimal patterns and speed of movements. Each of these categories is discussed separately, dealing with both the theoretical developments and the data that permit tests of the predictions. Studies that either develop testable predictions or that attempt to test such predictions in a precise quantitative manner are emphasized. The simple models so far formulated are supported reasonably well by available data. However, these simple models will require much modification, especially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated than just energy.