Cooperation invites cheating, and nowhere is this more apparent than when different species cooperate, known as mutualism. In almost all mutualisms studied, specialist parasites have been identified that purloin the benefits that one mutualist provides another. Explaining how parasites are kept from driving mutualisms extinct remains an unsolved problem because existing theories explaining the maintenance of cooperation do not apply to parasites of mutualisms. Nonetheless, these theories can be summarized in such a way as to suggest how mutualisms can persist in the face of parasites. (1) For cooperation to occur, the recipient of a benefit must reciprocate, and the recriprocated benefit must be captured by the initial giver or its offspring. (2) For cooperation to persist, the mutualism must be re-assembled each generation. Because most mutualisms are of the 'by-product' type, broadly defined, the first condition is normally always fulfilled. Thus, the maintenance of mutualism usually requires enforcement of the second condition: reliable re-assembly. Hence, I argue that the persistence of mutualism is best understood by using theories of species coexistence, because each mutualist can be considered a resource for the other, and species coexistence theory explains how multiple taxa (e.g. parasites and mutualists) can stably partition a resource over multiple generations. This approach connects the study of mutualism to theories of population regulation and helps to identify key factors that have promoted the evolution, maintenance and breakdown of mutualism. I discuss how these ideas might apply to and be tested in ant-plant, fig-wasp and yucca-moth mutualisms. Addicott JF'. 1996. Cheaters in yuccaJmoth mutualism. Nature 380 114-116. Addicott JF. 1986. Variation in the costs and benefits of mutualism: interaction between yuccas and yucca moths. Oecologca 70: 486494. Addicott JF. 1998. Regulation of mutualism between yuccas and yucca moths: population level processes. Oikos 81: 119-129. Addicott JF, Bao T. 1999. Limiting the costs of mutualism: multiple modes of interaction between yuccas and yucca moths. Proceedings of the Royal Society of London Series Addicott JF, Bronstein J, Kjellberg F. 1990. Evolution of mutualistic life cycles: yucca moths and fig wasps. In: Gilbert F, ed. Genetics, evolution, and coordination of insect life cycles. London: Springer, 143-161. Addicott JF, Tyre AJ. 1995. Cheating in an obligate mutualism: how often do yucca moths benefit yuccas? Oikos Aker CL, Udovic D. 1981. Oviposition and pollination behaviour of the yucca moth, Tegeticula maculata (Lepidoptera: Prodoxidae), and its relation to the reproductive biology of Yucca whipplei (Agavaceae). Oecologia 4 9 96-101. Anstett M-C, Bronstein JL, Hossaert-McKey M. 1996. Resource allocation: a conflict in the figfig wasp mutualism? Journal of Evolutionary Biology 9 417428. Armstrong RA. 1976. Fugitive species: experiments with fungi and some theoretical considerations. Ecology 57: 953-963. Axelrod R, DAmbrosio L. 1994. School of Public ...