In 2005, Sandi Willows-Munro (of the University of KwaZulu-Natal) and colleagues carried out a mitochondrial analysis of the nine "Tragelaphus" species. mtDNA and nDNA data were compared. The results showed that the tribe Tragelaphini is monophyletic with the lesser kudu basal in the phylogeny, followed by the nyala ("T. angasii"). On the basis of mitochondrial data, the lesser kudu separated from its sister clade around 13.7 million years ago. However, the nuclear data show that lesser kudu and nyala form a clade, and collectively separated from the sister clade 13.8 million years ago.