Solanum section Petota, which includes the potato and its wild relatives, is a difficult taxonomic group complicated by interspecific hybridization, introgression, allopolyploidy, a mixture of sexual and asexual reproduction, and possible recent species divergence (Spooner and van den Berg, 1992a;Spooner, 2009). Every taxonomist who has worked on the section has provided different hypotheses of the number of species, their interrelationships, and the hybrid origins of various taxa. A recent taxonomic conspectus (Spooner et al., 2014) recognized 107 species in section Petota, less than half as many as a treatment by Hawkes (1990) that recognized 232 species.Plastid DNA restriction site studies (Spooner et al., 1991;Spooner and Sytsma, 1992;RodrĂguez and Spooner, 1997;Spooner and Castillo, 1997) recognized four main clades in section Petota, referred to as clades 1-4. Subsequent studies using nuclear orthologs (Spooner et al., 2008b;RodrĂguez and Spooner, 2009;Cai et al., 2012) recovered three main clades, with similar interspecific relationships except they combined clades 1 and 2 with the nuclear clades then referred to as 1+2, 3, and 4. In-depth studies of these individual clades have been conducted with morphological and molecular data in clade 1+2 (RodrĂguez and Spooner, 1997; LaraCabrera and Spooner, 2004, 2005) and clade 3 (Ames et al., 2008;Ames and Spooner, 2010).Putatively related species complexes within the cultivated potato clade (clade 4) have been studied with DNA markers and morphological data, but never with a broad-scale phylogenetic study using multiple nuclear orthologs. According to the taxonomy of Spooner et al. (2014), the cultivated potato clade contains 34 predominately diploid species (some with rare triploid or tetraploid populations), two exclusively tetraploid species, and one species with diploid, tetraploid, and hexaploid populations. An additional 17 allopolyploid species share alleles with clades 1 or 3 and clade 4 (the cultivated potato clade) and are not studied here. The best-studied species complex in
PREMISE OF THE STUDY:The species boundaries of wild and cultivated potatoes are controversial, with most of the taxonomic problems in the cultivated potato clade. We here provide the first in-depth phylogenetic study of the cultivated potato clade to explore possible causes of these problems.
METHODS:We examined 131 diploid accessions, using 12 nuclear orthologs, producing an aligned data set of 14,072 DNA characters, 2171 of which are parsimony-informative. We analyzed the data to produce phylogenies and perform concordance analysis and goodness-of-fit tests.
KEY RESULTS:There is good phylogenetic structure in clades traditionally referred to as clade 1+2 (North and Central American diploid potatoes exclusive of Solanum verrucosum), clade 3, and a newly discovered basal clade, but drastically reduced phylogenetic structure in clade 4, the cultivated potato clade. The results highlight a clade of species in South America not shown before, 'neocardenasii' , sister to clade 1+2, t...