Should models of disease dynamics in herbivorous insects include the effects of variability in host-plant foliage quality?

Interactions between insects and their baculovirus pathogens are often described using simple disease models. Baculoviruses, however, are transmitted when insects consume virus-contaminated foliage, and foliage variability, whether within or between host-plant species, can affect viral infectiousness. Insect-baculovirus interactions may thus be embedded in a tritrophic interaction with the insect's host plant, but disease models include only the host and the pathogen. We tested these models by measuring the transmission of a baculovirus of gypsy moths (Lymantria dispar) on red oak (Quercus rubra) and white oak (Quercus alba) in the field in six experiments over four years. In all experiments, there were only weak effects of host-tree species, and in only one did the best-fitting model include tree species effects. These weak effects of foliage variability on transmission were not due to a lack of foliage variability on viral infectiousness, because when larvae were force-fed virus-contaminated foliage, infection rates were higher on white oak. Our results suggest that feeding behavior plays an important role in baculovirus transmission and that models can usefully describe baculovirus dynamics even without including foliage variability. Our work provides a clear example of how two-species models are sometimes sufficient to describe what appear to be tritrophic interactions.