White-tailed deer alter specialist and generalist insect herbivory through plant traits.

Within a plant species, leaf traits can vary across environmental, genetic, spatial, and temporal gradients, even showing drastic differences within individuals. Herbivory can also induce variation in leaf morphology, defensive structure, and chemistry including nutritional content. Indirect effects of prior insect herbivory on later herbivores have been well documented, but the induction of trait changes after vertebrate herbivory has been little explored. Here, we examined how browsing of spicebush (Lindera benzoin L.), a dominant understory shrub in eastern mesic forests, by white-tailed deer (Odocoileus virginianus L.) altered plant quality and subsequent foliar herbivory by insects. Browsing history explained ≈ 10% of overall leaf trait variation; regenerated leaves had greater water content and specific leaf area (P = 0.009), but were lower in nitrogen and greater in carbon (P < 0.001), than leaves on unbrowsed plants. However, browsing did not shift terpene chemistry as revealed by GC-MS. In the lab, caterpillars of the specialist spicebush swallowtail (Papilio troilus L.) preferred (P = 0.02) and grew 20% faster (P = 0.02) on foliage from browsed plants; whereas total herbivory in the field, including generalist insect herbivory, was twice as high on unbrowsed plants (P = 0.016). These results suggest that the ecological impacts of deer in forest understories can have cascading impacts on arthropod communities by changing the suitability of host-plants to insect herbivores.