Species Co-Occurrence Patterns among Lyme Borreliosis Pathogens in the Tick Vector Ixodes ricinus
ABSTRACTMixed infections have important consequences for the ecology and evolution of host-parasite interactions. In vector-borne diseases, interactions between pathogens occur in both the vertebrate host and the arthropod vector. Spirochete bacteria belonging to theBorrelia burgdorferisensu latogenospecies complex are transmitted byIxodesticks and cause Lyme borreliosis in humans. In Europe, there is a high diversity ofBorreliapathogens, and the main tick vector,Ixodes ricinus, is often infected with multipleBorreliagenospecies. In the present study, we characterized the pairwise interactions between fiveB. burgdorferisensu latogenospecies in a large data set ofI. ricinusticks collected from the same field site in Switzerland. We measured two types of pairwise interactions: (i) co-occurrence, whether double infections occurred more or less often than expected, and (ii) spirochete load additivity, whether the total spirochete load in double infections was greater or less than the sum of the single infections. Mixed infections ofBorreliagenospecies specialized on different vertebrate reservoir hosts occurred less frequently than expected (negative co-occurrence) and had joint spirochete loads that were lower than the additive expectation (inhibition). In contrast, mixed infections of genospecies that share the same reservoir hosts were more common than expected (positive co-occurrence) and had joint spirochete loads that were similar to or greater than the additive expectation (facilitation). Our study suggests that the vertebrate host plays an important role in structuring the community ofB. burgdorferisensu latogenospecies inside the tick vector.