Host plant limitation of butterflies in highly fragmented landscapes

Insect herbivores can be limited by host plants in two ways: density-dependent competition for food resources or density-independent search time limitation. Our understanding of density-dependent host plant limitation is relatively well developed and well integrated into conservation plans for at-risk insects. Search time limitation, a density-independent process, is much less well developed. Here, we explore both mechanisms using empirically based models of monarch butterfly population dynamics. These mechanisms differ fundamentally in their predictions: resource competition leads to matching of herbivore densities to host plant densities, and visible competition via consumption of host plants. Search time limitation leads to changes in population growth rate that can cause herbivore numbers to decline when host plant densities are constant. Search time limitation also implies that host plants can limit herbivores, even when many individual plants are uneaten. For monarch butterflies, our calculations suggest that many parts of North America have host plant densities below the threshold for search time limitation, which contrasts with the typical assumption of resource competition. More generally, incorporating search time limitation into conservation plans is important for reframing our understanding of how host plants limit insect herbivores in highly fragmented landscapes.