Late spring freezes coupled with warming winters alter temperate tree phenology and growth.

Spring phenology is advancing with warming but late spring freezes may not advance at the same rate, potentially leading to an increase in freezes after trees initiate budburst. Research suggests warming winters may delay budburst through reduced chilling, which may cause plants to leafout more slowly, thus decreasing spring freeze tolerance. Here, we assessed the effects of late spring freezes and reduced over-winter chilling on sapling phenology, growth and tissue traits, across eight temperate tree and shrub species in a laboratory experiment. We found that spring freezes delayed leafout - extending the period of greatest risk for freeze damage - increased damage to the shoot apical meristem, and decreased leaf toughness and leaf thickness. Longer chilling accelerated budburst and leafout, even under spring freeze conditions. Thus, chilling compensated for the adverse effects of late spring freezes on phenology. Despite the effects of spring freezes and chilling on phenology, we did not see any major reordering in the sequence of species leafout. Our results suggest climate change may impact forest communities not through temporal reassembly, but rather through impacts on phenology and growth from the coupled effects of late spring freezes and decreased over-winter chilling under climate change.