META-ANALYSIS OF CO-INFECTIONS IN TICKS

Microbial infections typically do not occur in isolation but co-occur within diverse communities of bacteria, fungi, protozoans, and viruses. Co-infections can lead to increased disease severity, lead to selection for increased virulence, and complicate disease diagnosis and treatment. Co-infections also occur in disease vectors, and represent one source of co-infections in hosts. We examined patterns of co-infections in ticks (Order Acari), which vector diverse human and wildlife pathogens, and asked whether the frequency of microbial co-infections deviated significantly from independent associations. Most published data were from Ixodes species and reported infection and co-infection frequencies of Borrelia burgdorferi and Anaplasma phagocytophilum. A total of 18 datasets representing 4978 adult ticks met our criteria for inclusion in the meta-analysis. Significant deviations from independent co-infection were detected in eight of the 18 populations. Five populations exhibited a significant excess of A. phagocytophilum/B. burgdorferi co-infections, including all populations of I. ricinus that deviated from independence. In contrast, both populations of I. persulcatus and one of two populations of I. scapularis exhibited a significant deficit of co-infection. The single population of I. pacificus examined had a significant excess of co-infection. Our meta-analyses indicate that tick-borne microbes are often distributed non-randomly, but the direction of deviation was not consistent, indicating that multiple mechanisms contribute to these patterns. Unfortunately, most published studies were not designed to describe patterns of co-infection, and provided insufficient data for our meta-analysis. Future studies should more explicitly measure and report co-infections in ticks, including co-infections by endosymbionts.