Facilitation drives tree seedling survival at alpine treelines

Alpine treelines are considered ecological monitors recording the impacts of climate change on trees and forests. To date, most treeline research has focused on how climate change drives treeline dynamics. However, little is known about how biotic interactions mediate treeline shifts, particularly in the case of tree recruitment, a bottleneck of treeline dynamics. We hypothesized that inter-and intraspecific facilitation determined the establishment and survival of tree seedlings at alpine treelines. To test this hypothesis, 630 Abies georgei var. smithii seedlings with different ages (4-6, 7-9 and 10-15 years old) were transplanted into three growth habitats (canopy-in, canopy-out and meadow) across the alpine treeline ecotone (4300-4500 m) in the Sygera Mountains, on the southeastern Qinghai-Tibetan Plateau. Microclimate, height growth, mortality rates and leaf functional traits of transplanted seedlings were measured over 3 years. We found that the variations in leaf functional traits were driven by microclimate. After the transplantation, the leaf concentrations of soluble sugars and starch and C:P ratio increased, whereas leaf size decreased. The resource use of seedlings gradually shifted to a more conservative strategy as indicated by changes in non-structural carbohydrates and nutrient concentrations. Radiation, temperature and moisture conditions, mediated by plant interactions, influenced seedling mortality and annual growth by affecting leaf morphological traits. Our findings illustrate how facilitation plays a crucial role in altering solar radiation and leaf trait functioning, determining seedling survival and growth at alpine treelines. We provide new insights into the underlying mechanisms for tree establishment and alpine treeline shifts in response to climate change.