The curse of the pharaoh hypothesis.

The 'curse of the pharaoh' has been used as a metaphor for the hypothesis that higher parasite propagule survival selects for higher virulence. Indeed, the mysterious death of Lord Carnavon after entering the tomb of the Egyptian pharaoh Tutankhamen could potentially be explained by an infection with a highly virulent and very long-lived pathogen. In this paper, I investigate whether parasite virulence increases with high propagule survival. In this respect, I derive an analytic expression of the evolutionarily stable level of parasite virulence as a function of propagule survival rate when the host-parasite system has reached a stable ecological equilibrium. This result shows that, if multiple infection occurs, higher propagule survival generally increases parasite virulence. This effect is enhanced when parasite dispersal coevolves with parasite virulence. In a more general perspective, the model shows the importance of taking into account the combination of direct and indirect effects (which I call inclusive effects) of higher transmission ability on the evolution of parasite virulence. The recognition of these effects has several practical implications for virulence management.