Reproductive phenology determines the linkages between radial growth, fruit production and climate in four Mediterranean tree species

The influence of climate on seed or fruit production and tree growth is a central question in forest ecology, with a key role on forest dynamics. However, the mechanisms linking inter- and intra-annual climate variability, fruiting and growth remain poorly understood, although they seem to be largely species-specific. The resource-matching hypothesis posits that fruit production responds to annual environmental variability, predicting a positive relationship between reproduction and growth, whereas other hypotheses such as resource switching or resource budget imply allocation trade-offs between reproduction and growth. Climate control over fruiting is thought to be more likely in variable, limiting environments such as those existing in seasonally dry Mediterranean forests. However, the studies about these forests are much scarcer than on temperate ecosystems, probably due to the sparseness and shortness of fruit production records. Here we analysed the relationships between monthly climate variables, tree-ring widths and fruit production at stand level in four tree species under different climatic conditions in Spain (12-27 trees and 24-54 radii were sampled in each site). For the species from xeric sites such as Pinus pinea, Castanea sativa and Quercus ilex, tree-ring width positively correlated with fruit production, as envisaged by the resource-matching hypothesis. This association was driven by prior wet winter climate conditions, which enhanced growth. Some of the significant correlations between climate variables and fruit yield corresponded to key reproductive phenophases. On the other hand, for the species from mesic sites (Abies alba) no positive growth-fruit production correlation was found, likely due to the mismatch in the effect of climatic factors on growth and fruiting. These results support the hypothesis that climatic conditions, particularly a positive water balance, play a major role as a proximate cause of fruit production in seasonally dry Mediterranean forests and underscore the importance of species-specific reproductive phenology traits.