A critical review of twenty years' use of the resource-ratio theory.

A model of species interactions based on their use of shared resources was proposed in 1972 by Robert MacArthur and later expanded in an article (1980) and a book (1982) by David Tilman. This "resource-ratio theory" has been used to make a number of testable predictions about competition and community patterns. We reviewed 1,333 papers that cite Tilman's two publications to determine whether predictions of the resource-ratio theory have been adequately tested and to summarize their general conclusions. Most of the citations do not directly test the theory: only 26 studies provide well-designed tests of one or more predictions, resulting in 42 individual tests of predictions. Most of these tests were conducted in the laboratory or experimental microcosms and used primary producers in freshwater systems. Overall, the predictions of the resource-ratio theory were supported 75% of the time. One of the primary predictions of the model, that species dominance varies with the ratio of resource availabilities, was supported by 13 of 16 tests, but most other predictions have been insufficiently tested. We suggest that more experimental work in a variety of natural systems is seriously needed, especially studies designed to test predictions related to resource supply and consumption rates.