Abstract. Widespread outbreaks of mountain pine beetles (MPB) are occurring throughout the range of this native insect. Episodic outbreaks are a common occurrence in the beetles' primary host, lodgepole pine. Current outbreaks, however, are occurring in habitats where outbreaks either did not previously occur or were limited in scale. Herein, we address widespread, ongoing outbreaks in high-elevation, whitebark pine forests of the Greater Yellowstone Ecosystem, where, due to an inhospitable climate, past outbreaks were infrequent and short lived. We address the basic question: are these outbreaks truly unprecedented and a threat to ecosystem continuity? In order to evaluate this question we (1) present evidence that the current outbreak is outside the historic range of variability; (2) examine system resiliency to MPB disturbance based on adaptation to disturbance and host defenses to MPB attack; and (3) investigate the potential domain of attraction to large-scale MPB disturbance based on thermal developmental thresholds, spatial structure of forest types, and the confounding influence of an introduced pathogen. We conclude that the loss of dominant whitebark pine forests, and the ecological services they provide, is likely under continuing climate warming and that new research and strategies are needed to respond to the crisis facing whitebark pine.

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Abstract. An innovative aerial survey method called the Landscape Assessment System (LAS) was used to assess mountain pine beetle (MPB; Dendroctonus ponderosae)-caused mortality of whitebark pine (Pinus albicaulis) across the species distribution in the Greater Yellowstone Ecosystem (GYE; 894 774 ha). This large-scale implementation of the LAS method consisted of 8673 km of flight lines, along which 4653 geo-tagged, oblique aerial photos were captured at the catchment level (a subset of 12-digit USGS hydrologic units) and geographic information system (GIS) processed. The Mountain Pine Beetle-caused Mortality Rating System, a landscape-scale classification system designed specifically to measure the cumulative effects of recent and older MPB attacks on whitebark pine, was used to classify mortality with a rating from 0 to 6 based on the amount of red (recent attack) and gray (old attack) trees visible. The approach achieved a photo inventory of 79% of the GYE whitebark pine distribution. For the remaining 21%, mortality levels were estimated based on an interpolated surface. Results that combine the photo-inventoried and interpolated mortality indicate that nearly half (46%) of the GYE whitebark pine distribution showed severe mortality (3-4 or 5.3-5.4 rating), 36% showed moderate mortality (2-2.9 rating), 13% showed low mortality (1-1.9 rating), and 5% showed trace levels of mortality (0-0.9). These results reveal that the proliferation of MPB in the subalpine zone of the GYE due to climate warming has led to whitebark pine mortality that is more severe and widespread than indicated from either previous modeling research or USDA Forest Service Aerial Detection surveys. Sixteen of the 22 major mountain ranges of the GYE have experienced widespread moderate-to-severe mortality. The majority of catchments in the other six mountain ranges show low-to-moderate mortality. Refugia from MPB outbreaks, at least for now, also exist and correspond to locations that have colder microclimates. The spatially explicit mortality information produced by this project has helped forest managers develop and implement conservation strategies that include both preservation and restoration efforts. Future research aimed at documenting and quantifying the ecological impacts of widespread decline and collapse of this foundation and keystone species is warranted.

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