Tree-ring isotopic imprints on time series of reproductive effort indicate warming-induced co-limitation by sink and source processes in stone pine.

Increasing evidence indicates that tree growth processes, including reproduction, can be either sink- or source-limited, or simultaneously co-limited by sink and source, depending on the interplay between internal and environmental factors. We tested the hypothesis that the relative strengths of photosynthate supply and demand by stem growth and reproduction create variable competition for substrate that is imprinted in the tree-ring isotopes (C and O) of stone pine (Pinus pinea L.), a masting gymnosperm with large costs of reproduction, under warming-induced drought. Across five representative stands of the Spanish Northern Plateau, we also identified reproductive phases where weather drivers of cone yield (CY) have varied over a 60-year period (1960-2016). We found that these drivers gradually shifted from winter-spring conditions 3 years before seed rain (cone setting) to a combination of 3- and 1-year lagged effects (kernel filling). Additionally, we observed positive regional associations between carbon isotope discrimination (Δ13C) of the year of kernel filling and CY arising at the turn of this century, which progressively offset similarly positive relationships between Δ13C of the year of cone setting and CY found during the first half of the study period. Altogether, these results pinpoint the increasing dependence of reproduction on fresh assimilates and suggest sink and source co-limitation superseding the sink-limited functioning of reproduction dominant before 2000. Under climate warming, it could be expected that drier conditions reinforce the role of source limitation on reproduction and, hence, on regeneration, forest structure and economic profit of the nutlike seeds of the species.