Costs and benefits for phytophagous myrmecophiles: When ants are not always available

Costs and benefits in mutualistic associations between ants and phytophagous myrmecophiles are context dependent. We collected information from the literature on costs and benefits of myrmecophily in aphids, coccids, membracids and lycaenids. A key result of the literature survey is that investment in mutualism with ants entails costs paid not only when ants are present (direct costs) but also when they are absent (indirect costs). We incorporated such a trade-off in a model that investigates the fitness consequences of the decision of a potential myrmecophile whether or not to invest in cooperation with ants. The model shows that whether myrmecophily should be favoured depends on the rate of increase of the population, and, if there are indirect costs, on the frequency of habitats with ants. Both direct and indirect costs can limit or prevent the evolution of myrmecophily even when ants are abundant. To understand the patterns of associations in the field we therefore need to measure the benefits and costs of myrmecophily both in the presence and in the absence of ants.