Seedling establishment at the alpine tree line - Can there be too much winter protection?

Abstract:

Alpine and arctic tree line expansion relies on tree seedling survival above the tree line, where the environment is harsh and protection by snow during winter is essential. Above the tree line, bryophytes are dominant; they may act as thermal insulators but their insulating ability differs between species. Apart from these positive effects, both snow and bryophytes may have negative effects on seedlings via shortening of the growing season or competition, respectively. Snow depth and duration are expected to change due to climate change, leading in some places to more snow and in others to less. What is the role of bryophytes insulating properties for seedling establishment under changing winter conditions at the alpine tree line? We hypothesized that protecting effects of snow and bryophytes would be more important for seedling survival in harsh climate (high elevation) than in milder climate (low elevation) (interactions: bryophyte*elevation and snow*elevation) and that negative effects of less snow would be ameliorated by well-insulating bryophytes (interaction: bryophyte*snow). To test this, we transplanted cores of three bryophyte species of differing insulation capacity and bare soil (control) from the subarctic tree line (~600m asl.) to 700 and 350 m asl. We transplanted 10 seedlings of two common tree line tree species (Betula pubescens and Pinus sylvestris) into each core in late summer. Cores were subjected to one of three snow treatments: autumn and spring snow removal or addition, or no manipulation. After the winter we scored seedling survival. The snow treatments had different effects at the two elevations (elevation* snow: P<0.0001) whereas bryophytes did not (elevation*bryophyte: n.s). In the harsh climate, snow addition generally enhanced seedling survival. In contrast, at the milder climate site, snow addition only increased survival in the bare soil treatment but decreased survival of seedlings in the bryophyte cores (bryophyte*snow: P=0.053). Our data show that snow cover is the dominant, positive control on winter survival of tree seedlings above the current tree line, irrespective of the bryophyte cover. In a milder climate, however, the role of snow cover depends on the bryophyte cover, with negative effects of extra snow on seedling survival in the presence of bryophytes and vice versa.