Regeneration dynamics of a mixed Mediterranean pine forest in the absence of fire

The aim of this study is to determine the competing regeneration and expansion patterns of two co-occurring pine species (Pinus brutia, Pinus nigra ssp. pallasiana), in a transitional montane Mediterranean zone. We measured the regeneration density of all woody species in 102 randomly located stands along an altitudinal gradient on the island of Lesbos, Greece. Individuals of pines were assigned to different size classes. Topographic factors (altitude, aspect, and soil depth) and light availability (through hemispherical photographs) were measured for each stand. Statistical analyses were applied to explore the effect of each factor on recruitment density of the competing pine species, and to elucidate patterns of interaction. Canopy openness was the most important parameter controlling the recruitment of P. brutia, while the regeneration density of P. nigra was mainly related to canopy openness and heat load. An idiosyncratic response of the recruitment vigour of the two species was identified along gradients of shade and drought stress. The decline in P. nigra recruitment density with drought conditions underlines threats to its population maintenance even in the absence of fire. On the other hand P. brutia seems to be a stronger invader in transitional zones. The studied species could be considered typical representatives of the two most widely distributed pine functional types across the Mediterranean basin, and our results agree with the theoretical ability of such species to maintain and expand their populations.