Climatic drivers and ecological implications of variation in the time interval between leaf-out and flowering

Leaf-out and flowering in any given species have evolved to occur in a predetermined sequence, with the inter-stage time interval optimized to maximize plant fitness. Although warming-induced advances of both leaf-out and flowering are well documented, it remains unclear whether shifts in these phenological phases differ in magnitudes and whether changes have occurred in the length of the inter-stage intervals. Here, we present an extensive synthesis of warming effects on flower-leaf time intervals, using long-term (1963-2014) and in situ data consisting of 11,858 leaf-out and flowering records for 183 species across China. We found that the timing of both spring phenological events was generally advanced, indicating a dominant impact of forcing conditions compared with chilling. Stable time intervals between leaf-out and flowering prevailed for most of the time series despite increasing temperatures; however, some of the investigated cases featured significant changes in the time intervals. The latter could be explained by differences in the temperature sensitivity (S_T) between leaf and flower phenology. Greater S_T for flowering than for leaf-out caused flowering times to advance faster than leaf emergence. This shortened the inter-stage intervals in leaf-first species and lengthened them in flower-first species. Variation in the time intervals between leaf-out and flowering events may have far-reaching ecological and evolutionary consequences, with implications for species fitness, intra/inter-species interactions, and ecosystem structure, function, and stability.