Numbers matter: how irruptive bark beetles initiate transition to self-sustaining behavior during landscape-altering outbreaks.

Irruptive forest insects such as bark beetles undergo intermittent outbreaks that cause landscape-scale tree mortality. Despite their enormous economic and ecological impacts, we still have only limited understanding of the dynamics by which populations transition from normally stable endemic to irruptive densities. We investigated density-dependent changes in mountain pine beetle reliance on stressed hosts, host selection, spatial configuration of attacks, and the interaction of host selection and spatial configuration by performing a complete census of lodgepole pine across six stands and 6 years. In addition, we compared the dynamics of mountain pine beetle with those of other bark beetles. We found that as population size increased, reliance on stressed trees decreased and new attacks shifted to larger trees with thicker phloem and higher growth rates that can support higher offspring production. Moreover, the spatial configuration of beetle-attacked trees shifted from random to spatially aggregated. Further, we found evidence that beetle utilization of larger trees was related to aggregation behavior as the size of tree attacked was positively correlated at 10-25 m, within the effective distance of pheromone-mediated signaling. In contrast, non-irruptive bark beetle species did not exhibit such density-dependent spatial aggregation at the stand scale or switches in host selection behavior. These results identify how density-dependent linkages between spatial configuration and host utilization can converge to drive population transitions from endemic to irruptive phases. Specifically, a combination of stand-level spatial aggregation, behavioral shifts, and higher quality of attainable hosts defines a critical threshold beyond which continual population growth becomes self-driving.