Abstract. In this study we present a simple optimization model for the evolution of defensive strategies (tolerance and resistance) of plants against their natural enemies. The model specifically evaluates the consequences of introducing variable costs and benefits of tolerance and resistance and nonlinear cost-and-benefit functions for tolerance and resistance. Incorporating these assumptions, the present model of plant defense predicts different evolutionary scenarios, not expected by previous work. Basically, the presence of an adaptive peak corresponding to intermediate levels of allocation to tolerance and resistance can arise when the shape parameter of the cost function is higher than the corresponding of the benefit function. The presence of two alternatives peaks of maximum tolerance and maximum resistance occurs only when benefits of tolerance and resistance interact less than additive. Finally, the presence of one peak of maximum resistance or maximum tolerance depends on the relative values of the magnitude of costs for tolerance and resistance. An important outcome of our model is that under a plausible set of conditions, variable costs of tolerance and resistance can represent an important aspect involved in the maintenance of intermediate levels of tolerance and resistance, and in favoring adaptive divergence in plant defensive strategies among populations. The model offers a framework for future theoretical and empirical work toward understanding spatial variation in levels of allocation to different defensive strategies.Key words. Evolutionarily stable strategy, host-pathogen interaction, plant-herbivore interaction, resistance, tolerance. During the last decade, the understanding of plant-enemy interactions has benefited by the incorporation into models of a ubiquitous type of defense (i.e., tolerance). Tolerance has been defined as the ability of a plant genotype to reduce the negative effects of consumers (e.g., herbivores, pathogens) on plant fitness (Rosenthal and Kotanen 1994;Strauss and Agrawal 1999;Stowe et al. 2000;Fornoni et al. 2003a). Unlike resistance (i.e., the ability of a plant to reduce the attack of natural enemies), tolerance is not believed to negatively affect the success of herbivore or pathogen populations (Rosenthal and Kotanen 1994;Fay et al. 1996; Roy and Kichner 2000;Tiffin 2000a;Restif and Koella 2003; but see Stinchcombe 2002). Consequently, the evolution of tolerance can limit antagonistic coevolution between plants and their enemies, whereas the evolution of resistance prolongs such coevolutionary outcome (Rausher 2001).The joint evolution of plant tolerance and resistance to natural enemies has attracted substantial theoretical attention over the last decade (Rosenthal and Kotanen 1994;Fineblum and Rausher 1995;Strauss and Agrawal 1999;Mauricio 2000;Roy and Kirchner 2000;Stowe et al. 2000;Tiffin 2000a;Restif andKoella 2003, 2004;Fornoni et al. 2003a). Several experimental studies have tested some of the predictions made by these theoretical analyses (Simms and T...