lesser kudu and the nyala are the basal most living species. Then around 10 million years ago there was a split between forest-dwelling, gracile species and larger, open-plains species. This renders Tragelaphus paraphyletic in respect to Taurotragus, which either means to make the latter genus into a junior synonym of the former genus or split Tragelaphus into several genera to maintain Taurotragus. In a 2011 large scale taxonomic reorganisation of ungulates by Groves and Grubb, they recognised new cryptic species, and split the traditional species into several geographical species. This was based on morphological comparisons between the species (in particular the dimensions of the skull elements and skin colouration) and defined species based on phylogenetic species concept. Groves and Grubb (2011) furthermore establish of several new genera in light of Tragelaphus being paraphyletic. They have a slight different arrangement of the species in Tragelaphus sensu stricto than suggested by genetics. Other scientists have mixed reactions to this proposal, with some preferring the traditional number of species to be seven and others preferring the new taxonomy from Groves and Grubb (2011). Below is the alternative taxonomy based on Groves and Grubb (2011), with species and subspecies names following Castelló (2016) from Bovids of the World: Tribe Tragelaphini Blyth, 1863 sensu Sokolov, 1953 – spiral-horned antelopes Genus Ammelaphus Heller, 1912 – lesser kudus Ammelaphus imberbis (Blyth, 1869) – northern lesser kudu Ammelaphus australis Heller, 1913 – southern lesser kudu Genus Nyala Heller, 1912 Nyala angasii (Angas, 1849) – lowland nyala Genus Taurotragus Wagner, 1855 – elands Taurotragus derbianus (Gray, 1847) – giant eland Taurotragus derbianus gigas Heuglin, 1863 – eastern giant eland Taurotragus derbianus derbianus (Gray, 1847) – western giant eland Taurotragus oryx (Pallas, 1767) – common eland Taurotragus oryx ivingstonii Sclater, 1864 – Livingstone's eland Taurotragus oryx pattersonianus Lydekker, 1906 – East African eland Taurotragus oryx oryx (Pallas, 1767) – Cape eland Genus Strepsiceros Hamilton Smith, 1827 – greater kudus Strepsiceros strepsiceros (Pallas, 1766) – Cape greater kudu Strepsiceros zambesiensis (Lorenz, 1894) – Zambezi greater kudu Strepsiceros chora (Cretzschmar, 1826) – northern greater kudu Strepsiceros cottoni (Dollman & Burlace, 1928) – western greater kudu Genus Tragelaphus de Blainville, 1816 Tragelaphus scriptus (Pallas, 1766) – western bushbuck Tragelaphus phaleratus (Hamilton Smith, 1827) – central bushbuck Tragelaphus bor Heuglin, 1877 – Nile bushbuck Tragelaphus decula (Rüppell, 1835) – Abyssinian bushbuck Tragelaphus meneliki Neumann, 1902 – Menelik's bushbuck Tragelaphus fasciatus Pocock, 1900 – eastern