Molecular phylogenetic studies in the Linaceae and Linum, with implications for their systematics and historical biogeography
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Best-known as the family of the cultivated flax, Linum usitatissimum L., the Linaceae is a small but ecologically diverse family of flowering plants, with approximately 250 species distributed throughout the temperate and tropical latitudes of the world. This work is an investigation of the systematics and biogeography of the family and a portion of its largest genus, Linum, using molecular phylogenetic methods. I collected DNA sequences of rbcL and matK genes from 51 species of Linaceae, representing all 14 genera, and combined them with data from 24 other families of the order Malpighiales in phylogenetic analyses. Results strongly support the monophyly of Linaceae and subfamily Linoideae in their current circumscriptions, but subfamily Hugonioideae is poorly supported. Molecular dating analysis suggests that the temperate Linoideae diversified in the Eocene or Oligocene, while tropical Hugonioideae diversified later, during the Miocene, perhaps ruling out Gondwanan vicariance as an explanation for their Pantropical distribution. Hugonia and Linum, the largest genera in their respective subfamilies, are each found have multiple segregate genera nested within them, indicating potential need for taxonomic revision of each subfamily. In Linoideae, I further investigate the phylogeny of a lineage that includes the yellow-flowered Linum sections Cathartolinum, Linopsis, and Syllinum, and the segregate genera Cliococca, Hesperolinon, Radiola, and Sclerolinon, to provide a framework in which to assess character evolution, classification, and biogeography. With data from four chloroplast markers (matK, ndhF, trnK 3’ intron, trnL-F region) and the nuclear ITS, and extensive sampling from Linum section Linopsis from Eurasia, Africa, and the Americas, Hesperolinon and Sclerolinon are shown to be related to a lineage of Central American linums including L. mexicanum Kunth and L. guatemalense Benth., while Cliococca is affiliated with South American Linum. The phylogeny supportes previous hypotheses of the evolution of some taxonomically important characters, and several well-supported lineages are identified which correspond to previously proposed taxonomic groupings. Results also provide evidence for a single trans-Atlantic disjunction and independent Old and New World colonizations of the southern hemisphere in yellow-flowered Linum, occurring during the Miocene.