Phylogenetic relationships among the "spiny solanums" (Solanum subgenus Leptostemonum, Solanaceae).
Species of Solanum subgenus Leptostemonum comprise almost one third of the genus and are distributed worldwide. Members of this group are defined by their sharp epidermal prickles; thus, they are commonly referred to as the "spiny solanums." This subgenus includes a number of economically important species such as the Old World eggplants, as well as locally cultivated New World species such as the naranjilla and cocona. Given the size and importance of this group we have examined phylogenetic relationships across subgenus Leptostemonum, including a large sampling of species from previously defined species groups within the subgenus. Evolutionary relationships were inferred using DNA sequence data from two nuclear regions (ITS and the granule-bound starch synthase gene [GBSSI or waxy]) and one chloroplast spacer region (trnS-trnG). Results suggest that Solanum subgenus Leptostemonum is monophyletic when the S. wendlandii and S. nemorense species groups are excluded. We have defined 10 clades within subgenus Leptostemonum, some of which correspond to previously circumscribed species groups or sections. Most of the Old World species of subgenus Leptostemonum belong to a single species-rich clade. Sharp prickles and/or stellate hairs evolved more than once in Solanum, and floral heterandry originated multiple times within the "spiny solanums".