1. Winter is a period of dormancy for plants of cold environments. However, winter climate is changing, leading to an increasing frequency of stochastic warm periods (winter warming events) and concomitant reductions in snow cover. These conditions can break dormancy for some plants and expose them to freeze-and-thaw stress. Mosses are a major component of high-latitude ecosystems, yet the longer-term impacts of such winter warming events on mosses remain unknown. 2. In order to determine the longer-term legacy effects of winter warming events on mosses, we undertook a simulation of these events over three consecutive winters in a sub-Arctic dwarf shrub-dominated open woodland. The mat-forming feather moss, Hylocomium splendens (the most abundant cryptogam in this system), is one of the most widespread Arctic and boreal mosses and plays a key functional role in ecosystems. We studied the ecophysiological performance of this moss during the summers of the experimental period (2007-2009) and in the following years (2010-2013). 3. We show that the previously reported warming-induced reduction in segment growth and photosynthesis during the experimental years was persistent. Four years after the last event, photosynthesis and segment growth were still 30 and 36% lower than control levels, which was only a slight improvement from 44 and 43% 4 years earlier. Winter warming did not affect segment symmetry. During the years after the last simulated event, in both warmed and control plots, chlorophyll fluorescence and segment growth, but not net photosynthesis, increased slightly. The increases were probably driven by increased summer rainfall over the study years, highlighting the sensitivity of this moss to rainfall change. 4. Overall, the legacy effects shown here demonstrate that this widespread and important moss is likely to be significantly disadvantaged in a future sub-Arctic climate where frequent winter warming events may become the norm. Given the key importance of mosses for soil insulation, shelter and carbon sequestration in high-latitude regions, such persistent impacts may ultimately affect important ecosystem functions.

Snow regimes affect biogeochemistry of boreal ecosystems and are altered by climate change. The effects on plant communities, however, are largely unexplored despite their influence on relevant processes. Here, the impact of snow cover on understory community composition and below-ground production in a boreal Picea abies forest was investigated using a long-term (8-year) snow cover manipulation experiment consisting of the treatments: snow removal, increased insulation (styrofoam pellets), and control. The snow removal treatment caused longer (118 vs. 57 days) and deeper soil frost (mean minimum temperature -5.5 vs. -2.2 degrees C) at 10 cm soil depth in comparison to control. Understory species composition was strongly altered by the snow cover manipulations; vegetation cover declined by more than 50% in the snow removal treatment. In particular, the dominant dwarf shrub Vaccinium myrtillus (-82%) and the most abundant mosses Pleurozium schreberi (-74%) and Dicranum scoparium (-60%) declined strongly. The C:N ratio in V. myrtillus leaves and plant available N in the soil indicated no altered nitrogen nutrition. Fine-root biomass in summer, however, was negatively affected by the reduced snow cover (-50%). Observed effects are attributed to direct frost damage of roots and/or shoots. Besides the obvious relevance of winter processes on plant ecology and distribution, we propose that shifts in the vegetation caused by frost damage may be an important driver of the reported alterations in biogeochemistry in response to altered snow cover. Understory plant performance clearly needs to be considered in the biogeochemistry of boreal systems in the face of climate change.