Compensatory Growth of Scots Pine Seedlings Mitigates Impacts of Multiple Droughts Within and Across Years

Tree seedling resistance to and recovery from abiotic stressors such as drought and warming are crucial for forest regeneration and persistence. Selection of more resilient provenances and their use in forest management programs might alleviate pressures of climate change on forest ecosystems. Scots pine forests in particular have suffered frequent drought-induced mortality, suggesting high vulnerability to extreme events. Here, we conducted an experiment using potted Scots pine seedlings from ten provenances of its south-western distribution range to investigate provenancespecific impacts of multiple drought events. Seedlings were grown under ambient and elevated temperatures for 1.5 years and were subjected to consecutive droughts during spring and summer. Growth (height, diameter, and needle) and spring phenology were monitored during the whole study period and complemented by biomass assessments (bud, needle, wood, and needle/wood ratio) as well as measurements of chlorophyll fluorescence and of needle stable carbon isotope ratio. Phenology, growth and biomass parameters as well as carbon isotope ratio and their (direct) responses to reoccurring droughts differed between provenances, indicating genotypic adaptation. Seedling growth was plastic during drought with intra-and inter-annual compensatory growth after drought stress release (carryover effects), however, not fully compensating the initial impact. For (smaller) seedlings from southern/drier origins, sometimes greater drought resistance was observed which diminished under warmer conditions in the greenhouse. Warming increased diameter growth and advanced phenological development, which was (partly) delayed by drought in 2013, but advanced in 2014. Earlier phenology was linked to higher growth in 2013, but interestingly later phenology had positive effects on wood and needle biomass when subjected to drought. Lastly, stable carbon isotope ratios indicated a clear drought response of carbon assimilation. Drought-induced reduction of the photosystem II efficiency was only observed under warmer conditions but showed compensation under ambient temperatures. Besides these direct drought impacts, also interactive effects of previous drought events were shown, either reinforcing or sometimes attenuating the actual impact. Thus, depending on amount and timing of events, Scots pine seedlings, particularly from southern origins, might be well adapted and resilient to drought stress and should be considered when discussing assisted migration under changing climatic conditions.