Untangling winter chilling and spring forcing effects on spring phenology of subtropical tree seedlings

Global warming is having an unprecedented impact on plant phenology and vitality worldwide, potentially leading to significant changes in the food web, carbon, water cycling and ecosystem functions. Environmental drivers explaining spring phenology, mainly including temperature (chilling in late autumn and winter and forcing in late winter and spring) and photoperiod, have been extensively investigated in temperate trees, but these factors have been rarely studied in subtropical forests and their potential effects are therefore relatively unknown. This knowledge gap is especially important to fill because these ecosystems harbor the largest portion of the world's biodiversity. In this study, we sought to test the effects of chilling (low vs. high) and forcing temperatures (20 vs. 25 degrees C) on spring phenology for seedlings of five subtropical woody species using six climate chambers. We compared forcing requirements for budburst and leaf-out in the different treatments by calculating the number of degree days achieved from the start of the experiment to the time of budburst and leaf-out, and we examined seedlings' survival under the different treatments. Although the survival of subtropical seedlings was found to be little affected by variation in chilling or forcing, longer chilling duration and warmer forcing tem-perature led to a lower forcing requirement for budburst, which advanced both budburst and leaf-out. This suggests that the seedlings experienced a non-linear accumulation of forcing with generally a higher efficiency at 25 degrees C than at 20 degrees C. Interestingly, shorter exposure to chilling conditions disrupted the sequence of budburst / leaf-out timings among the study species. This study confirms that the sensitivity of spring leaf phenology to forcing and chilling is not only found in temperate perennial plants but also in subtropical trees that undergo a dormancy period. It offers new perspectives for a comprehensive analysis of subtropical plant phenology in response to global climate change.