According to studies published in March 2015, odd-toed ungulates are in a close family relationship with at least some of the so-called Meridiungulata, a very diverse group of mammals living from the Paleocene to the Pleistocene in South America, whose systematic unity is largely unexplained. Some of these were classified on the basis of their paleogeographic distribution. However, a close relationship can be worked out to perissodactyls by means of protein sequencing and comparison with fossil collagen from remnants of phylogenetically young members of the Meridiungulata (specifically "Macrauchenia" from the Litopterna and "Toxodon" from the Notoungulata). Both kinship groups, the odd-toed ungulates and the Litopterna-Notoungulata, are now in the higher-level taxon of Panperissodactyla. This kinship group is included among the Euungulata which also contains the even-toed ungulates and whales (Artiodactyla). The separation of the Litopterna-Notoungulata group from the perissodactyls probably took place before the Cretaceous–Paleogene extinction event. "Condylarths" can probably be considered the starting point for the development of the two groups, as they represent a heterogeneous group of primitive ungulates that mainly inhabited the northern hemisphere in the Paleogene.