Efficacy of tree defense physiology varies with bark beetle population density: a basis for positive feedback in eruptive species

We evaluated the ability of constitutive and inducible defenses to protect trees and restrict herbivore reproduction across the endemic, incipient (i.e., transitory), and eruptive phases of a native bark beetle species. Host defenses were major constraints when mountain pine beetle (Dendroctonus ponderosae Hopkins) populations were low, but inconsequential after stand-level densities surpassed a critical threshold. We annually examined all lodgepole pines (Pinus contorta Douglas var. latifolia) in six 12-18 ha stands for 3-6 years for beetle attack and establishment as beetle densities progressed through various population phases. We also assayed a suite of tree physiological and chemical attributes and related them to subsequent attacks during that year. Rapidly inducible defenses appeared more important than constitutive defenses, and total monoterpenes were more important than particular constituents. Trees that exude more resin and accumulate higher monoterpene concentrations in response to simulated attack largely escaped natural attacks when populations were low. In stands where beetles had reached incipient densities, these defenses were ineffective. Larger diameter trees had more pronounced defenses than smaller diameter trees. As populations increased, beetles selected increasingly larger, more resource-rich trees, despite their better defenses. When populations were too low for cooperative attack, beetles exploited trees weakened by lower-stem insects. Behavioral plasticity allows beetles to persist at endemic levels until conditions shift, after which positive feedbacks predominate.