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Introgression frequently occurs between closely related plant species when there is incomplete reproductive isolation. However, few studies have been designed to examine the extent and direction of introgression from all three genomes, especially when they have different rates of gene flow. Conifers, in which chloroplast (cp), mitochondrial (mt) and nuclear (nr) genomes show contrasting rates of gene flow, provide a good model in which to carry out such a case study. Here we use data on sequence variation in 18 DNAs from three genomes for 311 individuals of 31 populations to investigate the extent and direction of introgressions between three spruce species in areas where they are sympatrically distributed. We found that one species seem to have introgressions derived from the other two species in areas of sympatric distribution; there were more introgressions from the maternally inherited mtDNA, which has the lowest rate of gene flow, and fewer introgressions from the bi-parentally inherited nrDNA. Further coalescent analysis of nrDNA population genetic data suggested that the species containing introgressions had experienced range expansion in the recent past while there was no such indication for the other two species. Our results support the hypotheses that introgressions occurred more frequently at genetic markers with lower rates of gene flow and that they usually took place from local species towards invading species that had undergone recent demographic expansion. [ABSTRACT FROM AUTHOR]

Elucidating the evolutionary history of current species diversity, especially trees with large effective population sizes and long generation times, is a complicated exercise confounded by gene flow and incomplete lineage sorting. In the present study, we aim to determine the origin and speciation of Picea schrenkiana and Picea smithiana using population genetic data from chloroplast (cp), mitochondrial (mt), and nuclear (nr) genomes. These two species occur in the Central Asian Highlands and Himalayas, respectively, where they are isolated from other Asian congeneric species by the Qinghai-Tibet Plateau (QTP) or adjacent deserts. Previous studies based on both morphological and molecular evidence suggest that they have contrasting phylogenetic relationships with Picea likiangensis or Picea wilsonii which are closely related and both located in the QTP. We examined genetic variation among 16 loci of three genomes from 30 populations of these four species. At both cpDNA loci and mtDNA loci, P. schrenkiana appeared to be closely related to P. likiangensis, although statistical support for this was weak. However, phylogenetic analyses and speciation tests based on the nuclear data from 11 loci provided evidence that P. schrenkiana and P. smithiana are sister species. These two species diverged around five million years ago (Mya) while the divergence between them and the P. likiangensis- P. wilsonii clade occurred about 18.4 Mya. We also detected gene flow accompanying these speciation events. Our results highlight the complex speciation histories of these alpine conifers due to interspecific gene flow and/or incomplete lineage sorting, and the importance of the early QTP uplifts in promoting the origin of these important conifer species in the Asian highlands. [ABSTRACT FROM AUTHOR]