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After the last glacial maximum the Alps have represented a major obstacle to the recolonisation of central and northern Europe by flora and fauna surviving in southern refugia. It is also believed that the Alps may have acted as a temporary refugia for some species, harbouring significant genetic variation or allowing the evolution of new genotypes and subspecies. To examine this possibility for white oak species, a total of 1375 samples from 292 locations were sampled in alpine and flanking regions of Austria, France, Germany, Italy and Switzerland and were assessed for cpDNA variation. Of the oak samples taken, throughout the entire alpine area Quercus robur was found most frequently. Quercus pubescens, widespread in Switzerland, Italy and up to the northern part of the French Alps was found south of 47°N and thus absent from Germany and sparse in Austria. Quercus petraea was mainly found in the parts of the Alps west of 12°E. Absent in the high elevations of the Austrian Alps Q. petraea and subspecies can be found at the eastern flank of the Alps. A total of 11 different haplotypes were found within the alpine area. Most frequent are different members of lineage A (common in the Balkan refugial area) with haplotype 7 found in 64% of the sample set. Haplotype 5, a more eastern member of the lineage, was found at the eastern flank of the Alps. Approximately 22% of the samples belonged to lineage C (from the Italian refuge), almost all of them were haplotype 1 with the majority crossing the Alps in Switzerland at the Simplon pass. Haplotype 2, the more eastern member of lineage C originating in Italy, is found at low frequency to the east of the Alps and in the Danube valley. Oak from refugial areas in Spain (lineage B) had only a limited impact on the alpine region. In spite of the historical anthropogenic influence in the alpine region, clear tracks of recolonisation were uncovered. In particular, the path of northward migration of oaks from Italy and the major route north of the Alps were clearly elucidated, confirming earlier assessment of pollen core data. However, the direction of migration (east to west or west to east) of oaks from the Balkan (haplotype 7) around the Alps still needs to be resolved.

A consortium of 16 laboratories have studied chloroplast DNA (cpDNA) variation in European white oaks. A common strategy for molecular screening, based on restriction analysis of four PCR-amplified cpDNA fragments, was used to allow comparison among the different laboratories. A total of 2613 oak populations (12, 214 individual trees from eight species) were sampled from 37 countries, and analysed with the four fragments. They belong to eight related oak species: Quercus robur, Q. petraea, Q. pubescens, Q. frainetto, Q. faginea, Q. pyrenaica, Q. canariensis and Q. macranthera. During this survey, 45 chloroplast variants were detected and are described together with their phylogenetic relationships, but several of these haplotypes were pooled when there were some risks of confusion across laboratories during the survey, and finally 32 remained that were mapped and used in diversity analyses. A strong phylogeographic structure is apparent from the data, where related haplotypes have broadly similar geographic distributions. In total, six cpDNA lineages are identified, which have distinct geographic distributions, mainly along a longitudinal gradient. Most haplotypes found in northern Europe are also present in the south, whereas the converse is not true, suggesting that the majority of mutations observed were generated prior to postglacial recolonisation, corroborating the conclusions of earlier studies. The description of a new western European lineage constitutes a major finding, compared to earlier phylogenetic treatments. Although the eight oak species studied systematically share cpDNA variants when in sympatry, they partition cpDNA diversity differently, as a consequence of their different ecology and life history attributes. Regional differences in levels of differentiation also exist (either species-specific or general) ; these seem to be related to the intensity of past and present management of the forests across Europe but also to the level of fragmentation of the range within these regions.

A consortium of 16 laboratories have studied chloroplast DNA (cpDNA) variation in European white oaks. A common strategy for molecular screening, based on restriction analysis of four PCR-amplified cpDNA fragments, was used to allow comparison among the different laboratories. A total of 2613 oak populations (12,214 individual trees from eight species) were sampled from 37 countries, and analysed with the four fragments. They belong to eight related oak species: Quercus robur, Q. petraea, Q. pubescens, Q. frainetto, Q. faginea, Q. pyrenaica, Q. canariensis and Q. macranthera. During this survey, 45 chloroplast variants were detected and are described together with their phylogenetic relationships, but several of these haplotypes were pooled when there were some risks of confusion across laboratories during the survey, and finally 32 remained that were mapped and used in diversity analyses. A strong phylogeographic structure is apparent from the data, where related haplotypes have broadly similar geographic distributions. In total, six cpDNA lineages are identified, which have distinct geographic distributions, mainly along a longitudinal gradient. Most haplotypes found in northern Europe are also present in the south, whereas the converse is not true, suggesting that the majority of mutations observed were generated prior to postglacial recolonisation, corroborating the conclusions of earlier studies. The description of a new western European lineage constitutes a major finding, compared to earlier phylogenetic treatments. Although the eight oak species studied systematically share cpDNA variants when in sympatry, they partition cpDNA diversity differently, as a consequence of their different ecology and life history attributes. Regional differences in levels of differentiation also exist (either species-specific or general); these seem to be related to the intensity of past and present management of the forests across Europe but also to the level of fragmentation of the range within these regions.

International audience

International audience; The aim of this study was to compare genetic diversity within and among Quercus spp. populations based on two contrasting types of nuclear markers. Seven mixed stands of Quercus petraea and Quercus robur were analysed using six highly polymorphic and codominantly inherited microsatellite markers as well as 165 dominant AFLP markers. Genetic differentiation and genetic diversity within each population were assessed. The intra- and inter-locus variances were calculated, and the results were used to compare the genetic diversity between populations. Both classes of markers revealed similar results: the genetic diversity within population and the genetic differentiation among populations is greater in Q. petraea than in Q. robur. The genetic differentiation is generally higher when AFLP markers were used in comparison to microsatelli- tes. For AFLPs, the inter-locus variance is always much higher than the intra-locus variance, and explains why it was not possible to distinguish populations using this marker system. Finally, no significant positive correlation was found between the level of within-population diversity assessed with the two markers.

The geographic distribution throughout Europe of each of 32 chloroplast DNA variants belonging to eight white oak species sampled from 2613 populations is presented. Clear-cut geographic patterns were revealed by the survey. These distributions, together with the available palynological information, were used to infer colonisation routes out of the glacial period refugia. In western Europe in particular, movements out of the Iberian and the Italian Peninsulas can be clearly identified. Separate refugia are also present in eastern Balkans, whereas further west in this peninsula similarities with Italy were evident. Movements resulting in the exchange of haplotypes between refugia both during the present interglacial and probably also during earlier glacial cycles were therefore inferred. The consequences of these past exchanges is that phylogenetically divergent haplotypes have sometimes followed very similar colonisation routes, limiting somewhat the phylogeographic structure. Cases of geographic disjunction in the present-day distribution of haplotypes are also apparent and could have been induced by the existence of rapid climatic changes at the end of the glacial period (specifically the Younger Dryas cold period), which resulted in range restriction following an early warm period during which oak first expanded from its primary refugia. This cold phase was followed by a new period of expansion at the outset of the Holocene, involving in some cases `secondary' refugia. It is expected that these short climate oscillations would have led to a partial reshuffling of haplotype distribution. Early association between haplotypes and oak species are also suggested by the data, although extensive introgression among species has ultimately largely blurred the pattern. This implies that colonisation routes may have been initially constrained by the ecological characteristics of the species hosting each chloroplast variant. We suggest for instance that two oak species distributed in the north of the Iberian Peninsula (Quercus petraea and Q. pubescens) are recent post-glacial immigrants there. When considered together, conclusions on the location of glacial period refugia and the colonisation routes derived from molecular information and fossil pollen data appear to be both largely compatible and complementary.

The geographic distribution throughout Europe of each of 32 chloroplast DNA variants belonging to eight white oak species sampled from 2613 populations is presented. Clear-cut geographic patterns were revealed by the survey. These distributions, together with the available palynological information, were used to infer colonisation routes out of the glacial period refugia. In western Europe in particular, movements out of the Iberian and the Italian Peninsulas can be clearly identified. Separate refugia are also present in eastern Balkans, whereas further west in this peninsula similarities with Italy were evident. Movements resulting in the exchange of haplotypes between refugia both during the present interglacial and probably also during earlier glacial cycles were therefore inferred. The consequences of these past exchanges is that phylogenetically divergent haplotypes have sometimes followed very similar colonisation routes, limiting somewhat the phylogeographic structure. Cases of geographic disjunction in the present-day distribution of haplotypes are also apparent and could have been induced by the existence of rapid climatic changes at the end of the glacial period (specifically the Younger Dryas cold period), which resulted in range restriction following an early warm period during which oak first expanded from its primary refugia. This cold phase was followed by a new period of expansion at the outset of the Holocene, involving in some cases secondary' refugia. It is expected that these short climate oscillations would have led to a partial reshuffling of haplotype distribution. Early association between haplotypes and oak species are also suggested by the data, although extensive introgression among species has ultimately largely blurred the pattern. This implies that colonisation routes may have been initially constrained by the ecological characteristics of the species hosting each chloroplast variant. We suggest for instance that two oak species distributed in the north of the Iberian Peninsula (Quercus petraea and Q. pubescens) are recent post-glacial immigrants there. When considered together, conclusions on the location of glacial period refugia and the colonisation routes derived from molecular information and fossil pollen data appear to be both largely compatible and complementary.

A consortium of 16 laboratories have studied chloroplast DNA (cpDNA) variation in European white oaks. A common strategy for molecular screening, based on restriction analysis of four PCR-amplified cpDNA fragments, was used to allow comparison among the different laboratories. A total of 2613 oak populations (12,214 individual trees from eight species) were sampled from 37 countries, and analysed with the four fragments. They belong to eight related oak species: Quercus robur, Q. petraea, Q. pubescens, Q. frainetto, Q. faginea, Q. pyrenaica, Q. canariensis and Q. macranthera. During this survey, 45 chloroplast variants were detected and are described together with their phylogenetic relationships, but several of these haplotypes were pooled when there were some risks of confusion across laboratories during the survey, and finally 32 remained that were mapped and used in diversity analyses. A strong phylogeographic structure is apparent from the data, where related haplotypes have broadly similar geographic distributions. In total, six cpDNA lineages are identified, which have distinct geographic distributions, mainly along a longitudinal gradient. Most haplotypes found in northern Europe are also present in the south, whereas the converse is not true, suggesting that the majority of mutations observed were generated prior to postglacial recolonisation, corroborating the conclusions of earlier studies. The description of a new western European lineage constitutes a major finding, compared to earlier phylogenetic treatments. Although the eight oak species studied systematically share cpDNA variants when in sympatry, they partition cpDNA diversity differently, as a consequence of their different ecology and life history attributes. Regional differences in levels of differentiation also exist (either species-specific or general); these seem to be related to the intensity of past and present management of the forests across Europe but also to the level of fragmentation of the range within these regions.

International audience

A total of 1113 oak trees from 222 populations originating from eight countries (Austria, Bosnia-Herzegovina, Croatia, Hungary, Romania, Slovakia, Slovenia, Yugoslavia) were sampled in natural populations or in provenance tests. The sampled trees belong to four different species (Quercus robur, Quercus petraea, Quercus pubescens, Quercus frainetto) and to several putative subspecies. Variation at four chloroplast DNA (cpDNA) fragments was studied using restriction enzymes, resulting in the detection of 12 haplotypes. One haplotype was present in 36% of the trees, and six were found in 6– 17% of the trees. The haplotypes are shared extensively between species and subspecies. They belong to three different lineages (A, C and E) and are phylogeographically structured in the region investigated. Haplotypes of lineage E dominate to the east of the Carpathian mountains in Romania, whereas the Carpathian Basin seems to have been colonised along several different colonisation routes, from the Balkan peninsula but also from Italy. The data support the possible role of climatic instability during the late glacial period in shaping this complex geographic structure. The presence of several secondary refugia could be inferred in the region, which have played a major role in the second step of recolonisation, at the onset of the Holocene period.