Space and the persistence of male–killing endosymbionts in insect populations

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Abstract
Male–killing bacteria are bacteria that are transmitted vertically through the females of their insect hosts. They can distort the sex ratio of their hosts by killing infected male offspring. In nature, male–killing endosymbionts (male killers) often have a 100% efficient vertical transmission, and multiple male–killing bacteria infecting a single population are observed. We use different model formalisms to study these observations. In mean–field models a male killer with perfect transmission drives the host population to extinction, and coexistence between multiple male killers within one population is impossible; however, in spatially explicit models, both phenomena are readily observed. We show how the spatial pattern formation underlies these results. In the case of high transmission efficiencies, waves with a high density of male killers alternate with waves of mainly wild–type hosts. The male killers cause local extinction, but this creates an opportunity for uninfected hosts to re–invade these areas. Spatial pattern formation also creates an opportunity for two male killers to coexist within one population: different strains create spatial regions that are qualitatively different; these areas then serve as different niches, making coexistence possible.