Krassimir D. Naydenov

Glacial vicariance in Eurasia: mitochondrial DNA evidence from Scots pine for a complex heritage involving genetically distinct refugia at mid-northern latitudes and in Asia Minor

BackgroundAt the last glacial maximum, Fennoscandia was covered by an ice sheet while the tundra occupied most of the rest of northern Eurasia. More or less disjunct refugial populations of plants were dispersed in southern Europe, often trapped between mountain ranges and seas. Genetic and paleobotanical evidences indicate that these populations have contributed much to Holocene recolonization of more northern latitudes. Less supportive evidence has been found for the existence of glacial populations located closer to the ice margin. Scots pine (Pinus sylvestris L.) is a nordic conifer with a wide natural range covering much of Eurasia. Fractures in its extant genetic structure might be indicative of glacial vicariance and how different refugia contributed to the current distribution at the continental level. The population structure of Scots pine was investigated on much of its Eurasian natural range using maternally inherited mitochondrial DNA polymorphisms.ResultsA novel polymorphic region of the Scots pine mitochondrial genome has been identified, the intron 1 of nad7, with three variants caused by insertions-deletions. From 986 trees distributed among 54 populations, four distinct multi-locus mitochondrial haplotypes (mitotypes) were detected based on the three nad7 intron 1 haplotypes and two previously reported size variants for nad1 intron B/C. Population differentiation was high (GST = 0.657) and the distribution of the mitotypes was geographically highly structured, suggesting at least four genetically distinct ancestral lineages. A cosmopolitan lineage was widely distributed in much of Europe throughout eastern Asia. A previously reported lineage limited to the Iberian Peninsula was confirmed. A new geographically restricted lineage was found confined to Asia Minor. A new lineage was restricted to more northern latitudes in northeastern Europe and the Baltic region.ConclusionThe contribution of the various ancestral lineages to the current distribution of Scots pine was asymmetric and extant endemism reflected the presence of large geographic barriers to migration. The results suggest a complex biogeographical history with glacial refugia shared with temperate plant species in southern European Peninsulas and Asia Minor, and a genetically distinct glacial population located more North. These results confirm recent observations for cold tolerant species about the possible existence of refugial populations at mid-northern latitudes contributing significantly to the recolonization of northern Europe. Thus, Eurasian populations of nordic plant species might not be as genetically homogenous as assumed by simply considering them as offsets of glacial populations located in southern peninsulas. As such, they might have evolved distinctive genetic adaptations during glacial vicariance, worth evaluating and considering for conservation.

Range-wide genetic structure of maritime pine predates the last glacial maximum: evidence from nuclear DNA

Using nuclear simple sequence repeats (nuSSRs), we determined the genetic variability in the natural distribution range of maritime pine (Pinus pinaster) in the western Mediterranean region. We analysed the role of global and significant climatic fluctuations in driving the evolutionary diversification of this species. We attempted to determine the impact of the last glacial maximum (LGM) and human activity on genetic variation and to identify the effect of bottlenecks, admixing, migration, time to the most recent common ancestor (TMRCA), and recent splits. A total of 972 individuals were analysed. The sample represented 27 natural populations from the western Mediterranean region, which encompasses most of the natural range of P. pinaster. Using eight nuSSRs, we analysed genetic diversity indices for each population and group of populations. We also examined the interpopulation structure by the frequency and distance method and investigated genetic barriers, signals of historical demographic fluctuations, phylogeographic structure, admixing, rate of mutation, migration, as well as testing the hypothesis of isolation by distance (IBD). Both cluster analyses showed similar population genetic structure with three genetic barriers that divided the samples into four large groups. Intensive migration was only detected during the period of the last glacial maximum (LGM), which permitted the mutation rate of the markers used to be calculated. The majority of the population was found to exhibit signs of a recent bottleneck and its timing showed a clear northeast-southwest geographic distribution. A clearly defined phylogeographic structure (Nst > Gst and Rst > Gst ) under IBD was established, and showed the highest divergence between groups of populations separated by physical barriers, such as the Strait of Gibraltar, the Mediterranean Sea and the Pyrenees. The high level of intergroup genetic differentiation (ΦIS = 20.26) was attributed to a long historical isolation (which occurred before the last 18 000 years) between the principal maritime pine population groups that occurred due to physical barriers that limited pollen and seed transfer, combined with a minimal effective radius of distribution. The low level of genetic diversity among the populations was combined with genetic drift and a recent bottleneck during the period of human activity. Significant migration across barriers was due to spontaneous phenomena during the LGM, which had no significant impact on the genetic structure owing to its relatively short duration and the fragmented species. The phylogeographic structure under the assumption of IBD was well established for P. pinaster in each of the principal population groups.