The Atlantic-Mediterranean watershed, river basins and glacial history shape the genetic structure of Iberian poplars.

Recent phylogeographic studies have elucidated the effects of Pleistocene glaciations and of Pre-Pleistocene events on populations from glacial refuge areas. This study investigates those effects in riparian trees (Populus spp.), whose particular features may convey enhanced resistance to climate fluctuations. We analysed the phylogeographic structure of 44 white (Populus alba), 13 black (Populus nigra) and two grey (Populus x canescens) poplar populations in the Iberian Peninsula using plastid DNA microsatellites and sequences. We also assessed fine-scale spatial genetic structure and the extent of clonality in four white and one grey poplar populations using nuclear microsatellites and we determined quantitative genetic differentiation (Q(ST) ) for growth traits in white poplar. Black poplar displayed higher regional diversity and lower differentiation than white poplar, reflecting its higher cold-tolerance. The dependence of white poplar on phreatic water was evidenced by strong differentiation between the Atlantic and Mediterranean drainage basins and among river basins, and by weaker isolation by distance within than among river basins. Our results suggest confinement to the lower river courses during glacial periods and moderate interglacial gene exchange along coastlines. In northern Iberian river basins, white poplar had lower diversity, fewer private haplotypes and larger clonal assemblies than in southern basins, indicating a stronger effect of glaciations in the north. Despite strong genetic structure and frequent asexual propagation in white poplar, some growth traits displayed adaptive divergence between drainage and river basins (Q(ST) >F(ST)), highlighting the remarkable capacity of riparian tree populations to adapt to regional environmental conditions.

The Strait of Gibraltar as a major biogeographic barrier in Mediterranean conifers: a comparative phylogeographic survey.

The Strait of Gibraltar (SG) is reputed for being both a bridge and a geographic barrier to biological exchanges between Europe and Africa. Major genetic breaks associated with this strait have been identified in various taxa, but it is unknown whether these disjunctions have been produced simultaneously or by independent biogeographic processes. Here, the genetic structure of five conifers distributed on both sides of the SG was investigated using mitochondrial (nad1 b/c, nad5-1, nad5-4 and nad7-1) and chloroplast (Pt1254, Pt15169, Pt30204, Pt36480, Pt71936 and Pt87268) DNA markers. The distribution of genetic variation was partially congruent between types of markers within the same species. Across taxa, there was a significant overlapping between the SG and the genetic breaks detected, especially for the four Tertiary species surveyed (Abies pinsapo complex, Pinus nigra, Pinus pinaster and Taxus baccata). For most of these taxa, the divergence of populations across the SG could date back to long before the Pleistocene glaciations. However, their strongly different cpDNA G(ST) and R(ST) values point out that they have had dissimilar population histories, which might include contrasting amounts of pollen-driven gene flow since their initial establishment in the region. The fifth species, Pinus halepensis, was genetically depauperated and homogenous on both sides of the SG. A further analysis of nuclear DNA sequences with coalescent-based isolation with migration models suggests a Pleistocene divergence of P. halepensis populations across the SG, which is in sharp contrast with the pre-Pleistocene divergence dates obtained for P. pinaster. Altogether, these results indicate that the genetic breaks observed across this putative biogeographical barrier have been produced by independent evolutionary processes related to the biological history of each individual species instead of a common vicariant phenomenon.

Spatial genetic structure in continuous and fragmented populations of Pinus pinaster Aiton.

Habitat fragmentation, i.e., the reduction of populations into small isolated remnants, is expected to increase spatial genetic structure (SGS) in plant populations through nonrandom mating, lower population densities and potential aggregation of reproductive individuals. We investigated the effects of population size reduction and genetic isolation on SGS in maritime pine (Pinus pinaster Aiton) using a combined experimental and simulation approach. Maritime pine is a wind-pollinated conifer which has a scattered distribution in the Iberian Peninsula as a result of forest fires and habitat fragmentation. Five highly polymorphic nuclear microsatellites were genotyped in a total of 394 individuals from two population pairs from the Iberian Peninsula, formed by one continuous and one fragmented population each. In agreement with predictions, SGS was significant and stronger in fragments (Sp = 0.020 and Sp = 0.026) than in continuous populations, where significant SGS was detected for one population only (Sp = 0.010). Simulations suggested that under fat-tailed dispersal, small population size is a stronger determinant of SGS than genetic isolation, while under normal dispersal, genetic isolation has a stronger effect. SGS was always stronger in real populations than in simulations, except if unrealistically narrow dispersal and/or high variance of reproductive success were modelled (even when accounting for potential overestimation of SGS in real populations as a result of short-distance sampling). This suggests that factors such as nonrandom mating or selection not considered in the simulations were additionally operating on SGS in Iberian maritime pine populations.

Mating system and pollen gene flow in Mediterranean maritime pine.

Mating systems define the mode of gene transmission across generations, helping to determine the amount and distribution of genetic variation within and among populations of plant species. A hierarchical analysis of Mediterranean maritime pine mating system (61 mother trees from 24 plots, clustered in three populations) was used to identify factors affecting mating patterns and to fit pollen dispersal kernels. Levels of ovule and seed abortion, multi- and single-locus outcrossing rates and correlated paternity were estimated from progeny arrays and correlated with ecological stand variables and biometric tree measures. Pollen dispersal kernels were fitted using TwoGener and KinDist indirect methods and simulations were carried out to identify relevant factors affecting correlated paternity. Maritime pine showed high outcrossing rates (t(m) and t(s) approximately 0.96) and relatively low levels of correlated paternity [an r(p) of 0.018 (Ritland's estimate) or 0.048 (Hardy's estimate)], although higher than in other anemophilous tree species. Mating system parameters had high variation at the single-tree level (99-100%) but no stand or population effect was detected. At the single-tree level, outcrossing rates were correlated with tree (diameter and height) and crown size. In addition, correlated paternity showed a significant negative correlation with tree height, height to crown base and height to the largest crown width, probably reflecting the importance of the trees' 'ecological neighborhoods'. Indirectly estimated pollen dispersal kernels were very leptokurtic (exponential-power distributions with beta<<0.5), with mean dispersal distances from 78.4 to 174.4 m. Fitted dispersal kernels will be useful in building explicit simulation models that include dispersal functions, and which will contribute to current conservation and management programs for maritime pine. Nevertheless, the numerical simulations showed that restricted dispersal, male fertility and phenological overlap could only partially explain the observed levels of correlated paternity; so other factors may also be relevant for the management of this valuable forest tree species.