Genetic signals of ancient decline in Aleppo pine populations at the species’ southwestern margins in the Mediterranean Basin

Microsatellite markers were used to characterize the structure of genetic diversity in natural Moroccan Aleppo pine (Pinus halepensis Mill.) populations, the most southwesterly marginal populations of the species in the Mediterranean Basin. Twenty-two natural populations and one artificial population, located in four regions covering most of the natural range of P. halepensis in the country, were sampled. Across this range, towards the south and west (and towards high altitudes) the populations become increasingly discrete and discontinuous. The nuclear microsatellite marker analysis suggests that a large proportion of the Aleppo pines in Morocco have derived from a single genetic lineage, represented by a central group of 11 of the examined populations located in the High and Middle Atlas Mountains. In addition, two smaller groups, represented by the marginal southwestern High Atlas populations, and three still smaller north / northeastern groups of populations located in the Rif and northeast Middle Atlas Mountains, could be genetically distinguished. Further, coalescence analysis of historical demographic population patterns suggests that ancient bottlenecks occurred in all of the natural populations. However, the population differentiation and genetic diversity levels we found were good (Fst=15.47), presumably because of the species' good potential for long-distance dispersal of seeds and high invasive capacity, which appear to have maintained a state of stable near-equilibrium, meta-population dynamics since ancient times