Hexaprotodon

The name Hexaprotodon was often applied to the pygmy hippopotamus before its reclassification into the genus Choeropsis. The genus has been historically applied to numerous fossil hippopotamus species spanning Asia, Africa and Europe. The genus sensu lato, has been suggested to be paraphyletic with respect to both species of living hippopotamus.[2] The uncontroversial, core Asian members of the genus most closely related to the type species H. sivalensis first appeared around 6 millon years ago, during the latest Miocene and were widespread throughout South and Southeast Asia, with the oldest records coming from the Siwalik Hills.[4][5] The African species Hexaprotodon bruneti from the Early Pleistocene of Ethiopia may be closely related to the Asian Hexaprotodon species, and thus belong in the genus in its more narrow sense. If so, it likely originates from a migration from Asia.[2]

The Asian species of Hexaprotodon, like the living hippopotamus (Hippopotamus amphibius), but unlike the pygmy hippoptamus are thought to have had a semiaquatic ecology, with their skull shape greatly resembling that of H. amphibius, with elevated orbits that allowed them to see above water while submerged. This lifestyle likely evolved independently in both Hexaprotodon and the genus Hippopotamus. In comparison to Hippopotamus, the mandibular symphysis is much more robust, the canine processes do not extend laterally outwards, and the molar teeth are lower crowned. The more slender and less massive postcranial skeleton compared to H. amphibius also suggests that Hexaprotodon was less adapted to walking in mud.[6] Dental microwear suggests a grazing diet for Asian Hexaprotodon species, similar to H. amphibius.[7]

Hexaprotodon was largely extinct by the Late Middle Pleistocene in Southeast Asia[4] but survived in Sumatra into the early Late Pleistocene, with one tooth dated to around 70,000 years ago.[5] The last known populations survived on the Indian Subcontinent to the very end of the Pleistocene.[4]Hexaprotodon survived in India until the terminal Pleistocene, with the latest dates being around 16,467–15,660 cal years Before Present from bones found in the Narmada River valley in central India. Fossil evidence from a late-surviving Indian Hexaprotodon indicates that it lived during a catastrophic drought caused by the latest Heinrich event, leading to an extremely weak Indian monsoon. It is thought that these drought conditions led to a heavy habitat fragmentation due to Hexaprotodon depending on aquatic habitats, prompting an extinction vortex. Humans may have also facilitated the extinction by hunting the hippopotamuses during this vulnerable state, although no evidence of hippopotamus butchery is known from the Indian subcontinent.[4]

The genus Hexaprotodon contains the following species, all from Asia and Africa:

Life Restoration of Hexaprotodon sivalensis

Genus sensu lato:

• Hexaprotodon bruneti (Boisserie and White, 2004)
• Hexaprotodon crusafonti (Aguirre, 1963)
• Hexaprotodon hipponensis (Gaudry, 1867)
• Hexaprotodon imagunculus (Hopwood, 1926)
• Hexaprotodon iravaticus (Falconer and Cautley, 1847)
• Hexaprotodon karumensis (Coryndon, 1977)
• Hexaprotodon mingoz (Boisserie et al., 2003)
• Hexaprotodon namadicus (Falconer and Cautley, 1847 - possibly same as H. palaeindicus)
• Hexaprotodon palaeindicus (Falconer and Cautley, 1847)
• Hexaprotodon pantanellii (Joleaud, 1920)
• Hexaprotodon primaevus (Crusafont et al., 1964)
• Hexaprotodon protamphibius (Arambourg, 1944)
• Hexaprotodon siculus (Hooijer, 1946)
• Hexaprotodon sinhaleyus (Deraniyagala)
• Hexaprotodon sivajavanicus (Hooijer, 1950)
• Hexaprotodon sivalensis (Falconer and Cautley, 1836)
• Hexaprotodon dhokwazirensis Akhtar and Bakr, 1995

Genus sensu stricto:

• Hexaprotodon bruneti (Boisserie and White, 2004)
• Hexaprotodon coryndoni
• Hexaprotodon dhokwazirensis (Akhtar and Bakr, 1995)
• Hexaprotodon iravaticus (Falconer and Cautley, 1847)
• Hexaprotodon namadicus (Falconer and Cautley, 1847 - possibly same as H. palaeindicus)
• Hexaprotodon palaeindicus (Falconer and Cautley, 1847)
• Hexaprotodon sinhaleyus (Deraniyagala)
• Hexaprotodon sivajavanicus (Hooijer, 1950)
• Hexaprotodon sivalensis (Falconer and Cautley, 1836)

H. imagunculus