Whitebark pine (Pinus albicaulis) has the largest and most northerly distribution of any white pine (Subgenus Strobus) in North America, encompassing 18˝latitude and 21˝longitude in western mountains. Within this broad range, however, whitebark pine occurs within a narrow elevational zone, including upper subalpine and treeline forests, and functions generally as an important keystone and foundation species. In the Rocky Mountains, whitebark pine facilitates the development of krummholz conifer communities in the alpine-treeline ecotone (ATE), and thus potentially provides capacity for critical ecosystem services such as snow retention and soil stabilization. The invasive, exotic pathogen Cronartium ribicola, which causes white pine blister rust, now occurs nearly rangewide in whitebark pine communities, to their northern limits. Here, we synthesize data from 10 studies to document geographic variation in structure, conifer species, and understory plants in whitebark pine treeline communities, and examine the potential role of these communities in snow retention and regulating downstream flows. Whitebark pine mortality is predicted to alter treeline community composition, structure, and function. Whitebark pine losses in the ATE may also alter response to climate warming. Efforts to restore whitebark pine have thus far been limited to subalpine communities, particularly through planting seedlings with potential blister rust resistance. We discuss whether restoration strategies might be appropriate for treeline communities.

Aim Alpine habitats support unique biodiversity confined to high‐elevation areas in the current interglacial. Plant diversity in these habitats may respond to area, environment, connectivity and isolation, yet these factors have been rarely evaluated in concert. Here we investigate major determinants of regional species pools in alpine grasslands, and the responses of their constituent species groups. Location European mountains below 50° N. Time period Between 1928 and 2019. Major taxa studied Vascular plants. Methods We compiled species pools from alpine grasslands in 23 regions, including 794 alpine species and 2,094 non‐alpines. We used species–area relationships to test the influence of the extent of alpine areas on regional richness, and mixed‐effects models to compare the effects of 12 spatial and environmental predictors. Variation in species composition was addressed by generalized dissimilarity models and by a coefficient of dispersal direction to assess historical links among regions. Results Pool sizes were partially explained by current alpine areas, but the other predictors largely contributed to regional differences. The number of alpine species was influenced by area, calcareous bedrock, topographic heterogeneity and regional isolation, while non‐alpines responded better to connectivity and climate. Regional dissimilarity of alpine species was explained by isolation and precipitation, but non‐alpines only responded to isolation. Past dispersal routes were correlated with latitude, with alpine species showing stronger connections among regions. Main conclusions Besides area effects, edaphic, topographic and spatio‐temporal determinants are important to understand the organization of regional species pools in alpine habitats. The number of alpine species is especially linked to refugia and isolation, but their composition is explained by past dispersal and post‐glacial environmental filtering, while non‐alpines are generally influenced by regional floras. New research on the dynamics of alpine biodiversity should contextualize the determinants of regional species pools and the responses of species with different ecological profiles.

Tree recruitment is a spatially structured process that may undergo change over time because of variation in postdispersal processes. We examined seed pilferage, seed germination, and seedling survival in whitebark pine to determine whether 1) microsite type alters the initial spatial pattern of seed caches, 2) higher abiotic stress (i.e. higher elevations) exacerbates spatial distribution changes, and 3) these postdispersal processes are spatially clustered. At two study areas, we created a seed distribution pattern by burying seed caches in microsite types frequently used by whitebark pine's avian seed disperser (Clark's nutcracker) in upper subalpine forest and at treeline, the latter characterized by high abiotic environmental stress. We monitored caches for two years for pilferage, germination, and seedling survival. Odds of pilferage (both study areas), germination (northern study area), and survival (southern study area) were higher at treeline relative to subalpine forest. At the southern study area, we found higher odds of 1) pilferage near rocks and trees relative to no object in subalpine forest, 2) germination near rocks relative to trees within both elevation zones, and 3) seedling survival near rocks and trees relative to no object at treeline. No microsite effects were detected at the northern study area. Findings indicated that the microsite distribution of seed caches changes with seed/seedling stage. Higher odds of seedling survival near rocks and trees were observed at treeline, suggesting abiotic stress may limit safe site availability, thereby shifting the spatial distribution toward protective microsites. Higher odds of pilferage at treeline, however, suggest rodents may limit treeline recruitment. Further, odds of pilferage were higher near rocks and trees relative to no object in subalpine forest but did not differ among microsites at treeline, suggesting pilferage can modulate the spatial structure of regeneration, a finding supported by limited clustering of postdispersal processes.

Many forest species are adapted to long-interval, high-severity fires, but the intervals between severe fires are decreasing with changes in climate, land use, and biological invasions. Although the effects of changing fire regimes on some important recovery processes have previously been considered, the consequences for the dispersal of propagules (plant seeds and fungal spores) in forest communities have not. We characterize three mechanisms by which changing fire regimes disrupt propagule dispersal in mesic temperate, boreal, and high-elevation forests: reduced abundance and altered spatial distributions of propagule source populations, less effective dispersal of propagules by wind, and altered behavior of animal dispersers and propagule predators. We consider how disruptions to propagule dispersal may interact with other factors that are also influenced by fire regime change, potentially increasing risk of forest conversion. Finally, we highlight urgent research topics regarding how dispersal limitation may shape twenty-first century forest recovery after stand-replacing fire.

In the northern Rocky Mountains of the U.S. and Canada, whitebark pine (Pinus albicaulis Engelm.) is a functionally important species in treeline communities. The introduced fungal pathogen Cronartium ribicola, which causes white pine blister rust, has led to extensive whitebark pine mortality nearly rangewide. We examined four treeline communities within the Greater Yellowstone Ecosystem (GYE) to assess structure and composition, whitebark pine prevalence and functional role, differences in growing season mesoclimate among study areas, and blister rust infection incidence. We found that (1) whitebark pine frequently serves as the majority overall, solitary, and leeward tree island conifer; (2) the prevalence of different tree species in the windward position in tree islands, and thus their potential as tree island initiators, may be predicted from their relative abundance as solitary trees; and (3) white pine blister rust infection incidence ranged from 0.6% to 18.0% across study areas. White pine blister rust poses a threat to treeline development and structure and the provision of ecosystem services in the GYE. Increasing blister rust resistance in nearby subalpine whitebark pine communities through seedling planting or direct seeding projects should eventually result in higher levels of blister rust resistance in whitebark pine in treeline communities.