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Chloroplast DNA evidence for the roles of island colonization and extinction in Tolpis (Asteraceae: Lactuceae)¹

²Section of Integrative Biology and Institute of Cellular and Molecular Biology, University of Texas, Austin, Texas 78712 USA; ³Department of Biological Sciences, Florida International University, University Park, Miami, Florida 33199 USA, and The Research Center, The Fairchild Tropical Garden, 11935 Old Cutler Road, Miami, Florida 33156 USA; ⁴Jardín de Aclimatación de La Orotava, Calle Retama Número 2, Puerto de la Cruz, Tenerife, Canary Islands E-38400 Spain

Tolpis consists of 13 species native to Africa, Europe, and Macaronesia, with at least one species endemic to each of the four major archipelagos of the Azores, Madeira Islands, Canary Islands, and Cape Verde Islands. All but two of these species develop woody stems by maturity. Chloroplast DNA restriction site variation was analyzed for all species of Tolpis and four outgroups in order to understand the patterns of island colonization and evolution of woodiness in this genus. Parsimony analyses revealed a strongly supported monophyletic Tolpis. Within the genus, the following three well-supported groups were detected: all species from the Canary Islands and Cape Verde Islands, both Azorean species, and both continental species. The Canary Island/Cape Verde clade was sister to the two continental species, and the Azorean clade was sister to this group. The two Madeiran species of Tolpis occupied the basalmost positions within the genus. When biogeography was mapped onto this phylogeny, nine equally parsimonious reconstructions (five steps each) of dispersal history were detected, which fell into two groups: eight reconstructions implied that Tolpis colonized Madeira from the continent, followed by continental extinction and subsequent continental recolonization, while one reconstruction implied that Tolpis colonized Macaronesia four times. Two of the reconstructions involving continental extinction required the least amount of overall dispersal distance. The cpDNA phylogeny also suggests that woodiness arose in the common ancestor of all extant Tolpis, followed by two independent reversals to an herbaceous habit. Assuming that one of the eight reconstructions favoring continental extinction and recolonization is true, our results suggest that Tolpis may represent the first documented example of a woody plant group in Macaronesia that has recolonized the mainland in herbaceous form.

Key Words: Asteraceae; chloroplast DNA phylogeny • Lactuceae • long-distance dispersal • Macaronesia • restriction site variation • Tolpis • woodiness

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