Patterns and processes in parasite co-infection.

Co-infection of individual hosts by multiple parasite species is a pattern that is very commonly observed in natural populations. Understanding the processes that generate these patterns poses a challenge. For example, it is difficult to discern the relative roles of exposure and susceptibility in generating the mixture and density of parasites within hosts. Yet discern them we must, if we are to design and deliver successful medical interventions for co-infected populations. Here, we synthesise an emergent understanding of how processes operate and interact to generate patterns of co-infection. We consider within-host communities (or infracommunities) generally, that is including not only classical parasites but also the microbiota that are so abundant on mucosal surfaces and which are increasingly understood to be so influential on host biology. We focus on communities that include a helminth, but we expect similar inferences to pertain to other taxa. We suggest that, thanks to recent research at both the within-host (e.g. immunological) and between-host (e.g. epidemiological) scales, researchers are poised to reveal the processes that generate the observed distribution of parasite communities among hosts. Progress will be facilitated by using new technologies as well as statistical and experimental tools to test competing hypotheses about processes that might generate patterns in co-infection data. By understanding the multiple interactions that underlie patterns of co-infection, we will be able to understand and intelligently predict how a suite of co-infections (and thus the host that bears them) will together respond to medical interventions as well as other environmental changes. The challenge for us all is to become scholars of co-infections.