Who acquires infection from whom? Estimating herpesvirus transmission rates between wild rodent host groups.

To date, few studies of parasite epidemiology have investigated 'who acquires infection from whom' in wildlife populations. Nonetheless, identifying routes of disease transmission within a population, and determining the key groups of individuals that drive parasite transmission and maintenance, are fundamental to understanding disease dynamics. Gammaherpesviruses are a widespread group of DNA viruses that infect many vertebrate species, and murine gammaherpesviruses (i.e. MuHV-4) are a standard lab model for studying human herpesviruses, for which much about the pathology and immune response elicited to infection is well understood. However, despite this extensive research effort, primarily in the lab, the transmission route of murine gammaherpesviruses within their natural host populations is not well understood. Here, we aimed to understand wood mouse herpesvirus (WMHV) transmission, by fitting a series of population dynamic models to field data on wood mice naturally infected with WMHV and then estimating transmission parameters within and between demographic groups of the host population. Different models accounted for different combinations of host sex (male/female), age (subadult/adult) and transmission functions (density/frequency-dependent). We found that a density-dependent transmission model incorporating explicit sex groups fitted the data better than all other proposed models. Male-to-male transmission was the highest among all possible combinations of between- and within-sex transmission classes, suggesting that male behaviour is a key factor driving WMHV transmission. Our models also suggest that transmission between sexes, although important, wasn't symmetrical, with infected males playing a significant role in infecting naïve females but not vice versa. Overall this work shows the power of coupling population dynamic models with long-term field data to elucidate otherwise unobservable transmission processes in wild disease systems.