Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Cladus: Synapsida
Cladus: Eupelycosauria
Cladus: Sphenacodontia
Cladus: Sphenacodontoidea
Cladus: Therapsida
Cladus: Theriodontia
Cladus: Cynodontia
Cladus: Eucynodontia
Cladus: Probainognathia
Cladus: Prozostrodontia
Cladus: Mammaliaformes
Classis: Mammalia
Subclassis: Trechnotheria
Infraclassis: Zatheria
Supercohors: Theria
Cohors: Eutheria
Infraclassis: Placentalia
Cladus: Boreoeutheria
Superordo: Laurasiatheria
Cladus: Scrotifera
Cladus: Ferungulata
Cladus: Euungulata
Ordo: Artiodactyla
Cladus: Artiofabula
Cladus: Cetruminantia
Subordo: Whippomorpha
Familia: Hippopotamidae
Genus: †Hexaprotodon
Species: †H. garyam – †H. madagascariensis – †H. sivalensis
Name
Hexaprotodon Falconer & Cautley, 1836: 51 [subgenus]
References
Primary references
Falconer, H. & Cautley, P.T. 1836. Notes on the fossil hippopotamus of the Sivalik Hills. Asiatic Researches 19: 39–53. BHL Reference page.
Links
Hexaprotodon in Mammal Species of the World.
Wilson, Don E. & Reeder, DeeAnn M. (Editors) 2005. Mammal Species of the World – A Taxonomic and Geographic Reference. Third edition. ISBN 0-8018-8221-4.
Vernacular names
Hexaprotodon is an extinct genus of hippopotamid known from Asia and possibly Africa and Europe. The name Hexaprotodon means "six front teeth" as some of the fossil forms have three pairs of incisors.[2][3] The pygmy hippopotamus was historically placed in the genus, but today is generally placed in its own genus. The core Asian members of the genus ranged from the Indian subcontinent to Southeast Asia, and are thought to have had an aquatic ecology similar to that of the living common hippopotamus. The last members of the genus became extinct during the Late Pleistocene, perhaps as recently as 15,000 years ago.
Taxonomy
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 million years ago, during the latest Miocene and were widespread throughout South and Southeast Asia, with the oldest records coming from the Siwalik Hills of the northern Indian subcontinent.[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]
Description
The Asian species of Hexaprotodon, like the living hippopotamus (Hippopotamus amphibius), but unlike the pygmy hippopotamus 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]
Extinction
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] with among 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]
Species
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 coryndoni
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 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
References
Seth, S. (1993), New Perspectives in Anthropology, M.D. Publications, p. 333, ISBN 9788185880198
Boisserie, Jean-Renaud (2005). "The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis". Zoological Journal of the Linnean Society. 143: 1–26. doi:10.1111/j.1096-3642.2004.00138.x.
Hexaprotodon[permanent dead link], The Paleobiology Database, läst 19 mars 2013.
Jukar, Advait M.; Patnaik, Rajeev; Chauhan, Parth R.; Li, Hong-Chun; Lin, Jih-Pai (September 2019). "The youngest occurrence of Hexaprotodon Falconer and Cautley, 1836 (Hippopotamidae, Mammalia) from South Asia with a discussion on its extinction". Quaternary International. 528: 130–137. Bibcode:2019QuInt.528..130J. doi:10.1016/j.quaint.2019.01.005. S2CID 133765385.
Smith, Holly E.; Price, Gilbert J.; Duval, Mathieu; Westaway, Kira; Zaim, Jahdi; Rizal, Yan; Aswan; Puspaningrum, Mika Rizki; Trihascaryo, Agus; Stewart, Mathew; Louys, Julien (2021-11-30). "Taxonomy, taphonomy and chronology of the Pleistocene faunal assemblage at Ngalau Gupin cave, Sumatra". Quaternary International. Human Evolution in the Asia-Pacific Realm: Proceedings of the 1st Asia-Pacific Conference on Human Evolution. 603: 40–63. doi:10.1016/j.quaint.2021.05.005. hdl:10072/404701. ISSN 1040-6182.
PANDOLFI, LUCA; MARTINO, ROBERTA; ROOK, LORENZO; PIRAS, PAOLO (2020-01-09). "INVESTIGATING ECOLOGICAL AND PHYLOGENETIC CONSTRAINTS IN HIPPOPOTAMIDAE SKULL SHAPE". Rivista Italiana di Paleontologia e Stratigrafia. 126 (1): V. 126 N. 1 (2020). doi:10.13130/2039-4942/12730.
Patnaik, Rajeev (June 2015). "Diet and habitat changes among Siwalik herbivorous mammals in response to Neogene and Quaternary climate changes: An appraisal in the light of new data". Quaternary International. 371: 232–243. doi:10.1016/j.quaint.2014.11.025.
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