Alpine treeline ecotones are potential refugia for a montane pine species threatened by bark beetle outbreaks

Warming-induced mountain pine beetle (Dendroctonus ponderosae; MPB) outbreaks have caused extensive mortality of whitebark pine (Pinus albicaulis; WBP) throughout the species' range. In the highest mountains where WBP occur, they cross alpine treeline ecotones (ATEs) where growth forms transition from trees to shrub-like krummholz, some of which survived recent MPB outbreaks. This observation motivated the hypothesis that ATEs are refugia for WBP because krummholz growth forms escape MPB attack and have the potential to produce viable seed. To test this hypothesis, we surveyed WBP mortality along transects from the ATE edge (locally highest krummholz WBP) downslope into the forest and, to distinguish if survival mechanisms are unique to ATEs, across other forest ecotones (OFEs) from the edge of WBP occurrence into the forest. We replicated this design at 10 randomly selected sites in the U.S. Northern Rocky Mountains. We also surveyed reproduction in a subset of ATE sites. Mortality was nearly absent in upper ATEs (mean ± SE percent dead across all sites of 0.03% ± 0.03% 0-100 m from the edge and 14.1% ± 1.7% 100-500 m from the edge) but was above 20% along OFEs (21.4 ± 5.2% 0-100 m and 32.4 ± 2.7% 100-500 m from the edge). We observed lower reproduction in upper ATEs (16 ± 9.9 cones/ha and 12.9 ± 5.3 viable seeds/cone 0-100 m from the edge) compared to forests below (317.1 ± 64.4 cones/ha and 32.5 ± 2.5 viable seeds/cone 100-500 m from the edge). Uniquely high WBP survival supports the hypothesis that ATEs serve as refugia because krummholz growth forms escape MPB attack. However, low reproduction suggests ATE refugia function over longer time periods. Beyond our WBP system, we propose that plant populations in marginal environments are candidate refugia if distinct phenotypes result in reduced disturbance impacts.