Aggregation and a strong Allee effect in a cooperative outbreak insect.

Most species that are negatively impacted when their densities are low aggregate to minimize this effect. Aggregation has the potential to change how Allee effects are expressed at the population level. We studied the interplay between aggregation and Allee effects in the mountain pine beetle (Dendroctonus ponderosae Hopkins), an irruptive bark beetle that aggregates to overcome tree defenses. By cooperating to surpass a critical number of attacks per tree, the mountain pine beetle is able to breach host defenses, oviposit, and reproduce. Mountain pine beetles and Hymenopteran parasitoids share some biological features, the most notable of which is obligatory host death as a consequence of parasitoid attack and development. We developed spatiotemporal models of mountain pine beetle dynamics that were based on the Nicholson-Bailey framework but which featured beetle aggregation and a tree-level attack threshold. By fitting our models to data from a local mountain pine beetle outbreak, we demonstrate that due to aggregation, attack thresholds at the tree level can be overcome by a surprisingly low ratio of beetles per susceptible tree at the stand level. This results confirms the importance of considering aggregation in models of organisms that are subject to strong Allee effects.