Species climatic niche explains post-fire regeneration of Aleppo pine (Pinus halepensis Mill.) under compounded effects of fire and drought in east Spain

Fire and drought are two major agents that shape Mediterranean ecosystems, but their interacting effects on forest resilience have not been yet fully addressed. We used Pinus halepensis to investigate how compound fire-drought regimes determine the success of post-fire regeneration. We measured the density of P.halepensis newly established individuals following fire in forty-three sites along the Spanish east coast, the wetter region of the species distribution. The climatic niche of P.halepensis was characterized by considering their populations across its Spanish distribution range. We used yearly values (1979-2013 period) of accumulated precipitation, mean temperature and the warmest quarter values of these two variables to generate the climatic space or climatic niche occupied by the species. Kernel density estimates were then applied to determine the niche centroid, which would correspond to the species' climatic optimum within its Spanish distribution range. Then, we computed the pre- and post-fire climatic deviations of each sampling site as the difference between site-specific climate conditions respect to the species niche centroid, and assessed their relationship with the success of post-fire regeneration. We found highly variable patterns of post-fire regeneration density of P.halepensis over the studied sites, ranging from 7 to 42,822 tree pines ha^-1. Generalized linear models indicated a positive relationship between fire severity and the density of P.halepensis regeneration. Positive temperature deviations - warm conditions - before fire were positively related to pine regeneration. This effect increases under higher fire severity. By contrast, warm temperatures after fire showed a negative effect on the density of pine trees. Positive precipitation deviations - wet conditions - after fire enhanced pine regeneration, while precipitation before fire did not had any significant effect. Though P.halepensis is considered a species adapted to fire and drought, the interaction between these two disturbances can alter the success of its post-fire recovery patterns limiting the species' resilience in the future.