Defoliator outbreaks track with warming across the Pacific coastal temperate rainforest of North America

The biogeography of irruptive insect herbivores is determined by host availability and climate conditions. As such, outbreak distributions are sensitive to climatic change, especially across large latitudinal gradients. Here, we investigate the outbreak distributions of two understudied defoliators, hemlock sawfly Neodiprion tsugae (Hymenoptera) and western blackheaded budworm Acleris gloverana (Lepidoptera), that have both recently impacted the greatest land area recorded across the Pacific coastal temperate rainforest since the establishment of aerial survey programs. We compiled polygon-based estimates of insect damage collected by aerial observers, forest inventory, and downscaled climatic data to develop gridded estimates of bioclimatic conditions across the extent of the Pacific coastal temperate rainforest, including the continental United States, British Columbia and Alaska. We leveraged these data to develop ensemble machine learning models with the goal of predicting the outbreak distribution of each insect. In this manuscript we: 1) describe the historical patterns of defoliator outbreaks, 2) identify and describe climatic conditions associated with outbreaks in both species and 3) assess whether historic outbreaks have tracked geographic shifts in climate conditions across the region. We demonstrate that outbreaks of hemlock sawfly and western blackheaded budworm have been observed across the Pacific coastal temperature rainforests of North America in each decade since the establishment of the Canadian and United States aerial survey programs. The distribution of outbreaks by both insects were best explained by host availability, a limited range of spring, summer, and winter temperatures, and minimum precipitation. Finally, we demonstrate that outbreaks have tracked the poleward shift in suitable climate over the last century. This study establishes a baseline understanding of the climatic constraints and biogeographic patterns of historic sawfly and budworm outbreaks across the Pacific coastal temperate rainforest and emphasizes the overarching importance of climate in driving the irruptive dynamics of these defoliator species.

Keywords: climate envelope, defoliators, hemlock sawfly, Pacific coastal temperate rainforest, population dynamics, western blackheaded budworm