Boreal tree species are expected to invade tundra ecosystems as climate warms. Because forested ecosystems differ from tundra ecosystems in a number of climatically relevant characteristics, this advance of the altitudinal and latitudinal tree limit may ultimately feedback on regional climate. We used tree rings to reconstruct the response of the treeline ecotone on the Seward Peninsula in northwestern Alaska to 20th century warming. We further explored the sensitivity of the treeline ecotone to climate change using the spatially explicit, frame-based model ALFRESCO. Our reconstructions of forest response to past warming indicate that in upland tundra spruce have successfully established progressively farther from the forest limit since the 1880s. Shrub tundra has thus been converted to low-density forest-tundra within a band extending approximately 10 km from the forest limit. In lowland sites, where soil thermal and hydrologic properties are the primary constraint on tree distribution, trees began establishing in tundra after 1920, and establishment was restricted to areas that had experienced thermokarst activity in the past. Modeling experiments suggested that changes in disturbance regime and spruce growth response to climate may introduce strongly nonlinear responses to climate change at treeline. Field data and model experiments thus both indicate that large and nearly instantaneous responses to warming are likely at the treeline ecotone but that sensitivity to warming is likely to vary substantially over space and time.