Cross-scale drivers of natural disturbances prone to anthropogenic amplification: The dynamics of bark beetle eruptions

Biome-scale disturbances by eruptive herbivores provide valuable insights into species interactions, ecosystemfunction, and impacts of global change. We present a conceptual framework using one system as a model, emphasizing interactions across levels of biological hierarchy and spatiotemporal scales. Bark beetles are major natural disturbance agents in western North American forests. However, recent bark beetle population eruptions have exceeded the frequencies, impacts, and ranges documented during the previous 125 years. Extensive host abundance and susceptibility, concentrated beetle density, favorable weather, optimal symbiotic associations, and escape from natural enemies must occur jointly for beetles to surpass a series of thresholds and exert widespread disturbance. Opposing feedbacks determine qualitatively distinct outcomes at junctures at the biochemical through landscape levels. Eruptions occur when key thresholds are surpassed, prior constraints cease to exert influence, and positive feedbacks amplify across scales. These dynamics are bidirectional, as landscape features influence how lower-scale processes are amplified or buffered. Climate change and reduced habitat heterogeneity increase the likelihood that key thresholds will be exceeded, and may cause fundamental regime shifts. Systems in which endogenous feedbacks can dominate after external forces foster the initial breach of thresholds appear particularly sensitive to anthropogenic perturbations.