Phylogenetic placement of two enigmatic genera, Borthwickia and Stixis, based on molecular and pollen data, and the description of a new family of Brassicales, Borthwickiaceae

Capparaceae (Brassicales) as traditionally circumscribed is heterogeneous, and several genera have been segregated from it based on molecular and/or morphological data. However, Borthwickia and Stixis, two Southeast Asian endemic genera of Capparaceae with controversial positions, have not previously been evaluated in a molecular phylogenetic study. Here, we used four plastid DNA regions (matK, ndhF, rbcL, trnL-trnF) and pollen data to determine their phylogenetic relationships within core Brassicales. Our results showed that neither Borthwickia nor Stixis is a member of Capparaceae. The two genera, together with Forchhammeria, Gyrostemonaceae, Resedaceae, and Tirania, formed a clade with strong support. Stixis is closely related to Tirania, a relationship that is also supported by morphological characters, such as six sepals and three- or four-locular ovaries. Most interestingly, Borthwickia was resolved as sister to the Forchhammeria-Resedaceae-Stixis-Tirania clade with moderate to strong support. However, Borthwickia differs markedly from its sister group in having opposite leaves, one indistinct stigma, more than four carpels and locules, a linear ovary with ridges, and pollen grains with perforate exine sculpturing. Thus, we describe a new family, Borthwickiaceae, for the genus.