Insect herbivores, density dependence, and the performance of the perennial herb Solanum carolinense

How insect herbivores affect plant performance is of central importance to basic and applied ecology. A full understanding of herbivore effects on plant performance requires understanding interactions (if any) of herbivore effects with plant density and size because these interactions will be critical for determining how herbivores influence plant population size. However, few studies have considered these interactions, particularly over a wide enough range of densities to detect nonlinear effects. Here we ask whether plant density and herbivores influence plant performance linearly or nonlinearly, how plant density affects herbivore damage, and how herbivores alter density dependence in transitions between plant size classes. In a large field experiment, we manipulated the density of the herbaceous perennial plant Solanum carolinense and herbivore presence in a fully crossed design. We measured plant size, sexual reproduction, and damage to plants in two consecutive years, and asexual reproduction of new stems in the second year, allowing us to characterize both plant performance and rates of transition between plant size classes across years. We found nonlinear effects of plant density on damage. Damage by herbivores and plant density both influenced sexual and asexual reproduction of S. carolinense; these effects were mostly mediated via effects on plant size. Importantly, we found that herbivores altered the pattern of linear density dependence in some transition rates (including survival and asexual reproduction) between plant size classes. These results suggest that understanding the ecological or evolutionary effects of herbivores on plant populations requires consideration of plant density and plant size, because feedbacks between density, herbivores, and plant size may complicate longer-term dynamics.