Mafia behaviour and the evolution of facultative virulence.

Some organisms enforce "maladaptive" behaviours on others of the same or different species by imposing costs in the absence of compliance. Such enforcement is used by the enforcer to obtain benefits in the possession of the enforced individual. This mechanism is known as mafia behaviour in humans, but may be widespread in parasite-host relationships in nature, from the cellular level to societies. In this paper we describe the evolution of such mafia mechanisms, and we propose a fuzzy logic model where the mafia mechanism is based on enforcement of hosts by exponentially increasing the cost of resistance to the parasite. The benefits of host resistance can be counteracted by parasite virulence, or even a decrease in response to an increment in its resistance. This parasite response to the host defence increment can be used for the parasite to teach the host that it is better to pay part of its benefits than increase its extremely costly defence. This model differs from others because it takes into account the evolution of host defence related to the evolution of parasite virulence (host-parasite coevolution) and points out an optimum in host defence related to the facultative virulence of the parasite. We provide several potential examples of facultative virulence depending on the antiparasite responses of hosts, and we suggest that this kind of mafia behaviour may be a widespread mechanism in biological processes at a number of different levels.