Winter is coming: plant freezing resistance as a key functional trait for the assembly of annual Mediterranean communities.

Background and Aims: In Mediterranean annual plants, germination mainly occurs during the autumn and only those seedlings that survive winter freezing can flower and produce seedlings in spring. Surprisingly, the effect of freezing events as an abiotic determinant of these communities remains unexplored. The present study aimed to investigate how freezing events affect annual Mediterranean communities and whether their functional structure as related to freezing resistance is linked to the main biotic and abiotic determinants of these communities. Methods: In 120 plots located on a semi-arid Mediterranean steppe (Spain), the community functional structure related to the lethal temperature causing 50 % frost damage (LT50 trait) in seedlings was estimated and summarized as the community-weighted mean (CWM-LT50) and its functional diversity (FD-LT50). Plots were stratified according to distance to rabbit shelters and latrines as a proxy for rabbit density, proximity to Stipa tenacissima and spring water availability, where annual species abundance was recorded in all plots over three consecutive years. Key Results: Annual species were able to resist a threshold temperature of -4 °C and most had LT50 values around the absolute minimum temperature (-9.5 °C) in the three years. Higher rabbit densities led to lower CWM-LT50 and higher FD-LT50 values. Plots close to Stipa tussocks had higher CWM-LT50 values whereas water availability had no effects. Conclusions: High freezing resistance was extended among winter annual species, suggesting the presence of an association between historical environmental filtering and low winter temperatures. However, the community functional structure related to freezing resistance remained variable among scenarios with differences in herbivory pressure and distance to perennial vegetation. The trends observed indicate that traits that allow plants to deal with herbivory may also promote freezing resistance, and that tussocks can act as nurses via microclimatic amelioration of harsher winter conditions.