Northern Hemisphere disjunctions in Lactuca (Cichorieae, Asteraceae): independent Eurasia to North America migrations and allopolyploidization

The Lactuca lineage is one of nine lineages in the lettuce subtribe (Cichorieae, Asteraceae) distributed in Europe, Africa, Asia and North America. Within the Lactuca lineage two clades show disjunct Eurasian-North American distributions. One disjunct Glade consists of diploids (x = 8) and allotetraploids (x = 17), the former restricted to Eurasia and the latter to North America and the Azores. In contrast, members of the other Eurasian-North American disjunct Glade are all diploid (x = 9), like the remainder of the Lactuca lineage (diploid, x = 8 or 9). The aims of the present study were to investigate the migration pathways that led to the disjunct distributions of these two Eurasian-North American clades and the potential progenitors of the allopolyploid taxa. We conducted deep taxon sampling and multi-locus phylogenetic analyses using nuclear ribosomal DNA (ETS and ITS), a low-copy nuclear marker (A44) and five non-coding plastid markers. Divergence time estimations with BEAST and ancestral biogeographic estimations with BioGeoBEARS suggested that both lineages reached North America by the late Miocene. Cloning of the A44 region revealed two sequence copies within allopolyploid individuals that were resolved in divergent clades and this helped to identify potential progenitors. We provide competing hypotheses for the progenitor species and biogeographic pathways that gave rise to the allotetraploid lineage, and we propose a North American origin for the Azorean endemic. Taxonomic conclusions include L. graminifolia var. mexicana being raised to specific rank with the name L. brachyrrhyncha and the alleged endemic L. jamaicensis in fact represents the SE Asian L. indica, introduced to Jamaica.