Hexaprotodon
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''Hexaprotodon'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
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 Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
.''Hexaprotodon''
The Paleobiology Database, läst 19 mars 2013.
The
pygmy hippopotamus The pygmy hippopotamus or pygmy hippo (''Choeropsis liberiensis'') is a small Hippopotamidae, hippopotamid which is native to the forests and swamps of West Africa, primarily in Liberia, with small populations in Sierra Leone, Guinea, and Ivory Co ...
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 The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division ...
, perhaps as recently as 15,000 years ago. Extinct animals of Indonesia


Taxonomy

The name ''Hexaprotodon'' was often applied to the
pygmy hippopotamus The pygmy hippopotamus or pygmy hippo (''Choeropsis liberiensis'') is a small Hippopotamidae, hippopotamid which is native to the forests and swamps of West Africa, primarily in Liberia, with small populations in Sierra Leone, Guinea, and Ivory Co ...
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 ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'', has been suggested to be
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
with respect to both species of living hippopotamus. The uncontroversial, core Asian members of the genus most closely related to the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''H. sivalensis'' first appeared around 6 million years ago, during the latest
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
and were widespread throughout
South South is one of the cardinal directions or compass points. The direction is the opposite of north and is perpendicular to both west and east. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Proto-Germanic ''*sunþa ...
and
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
, with the oldest records coming from the
Siwalik Hills The Sivalik Hills, also known as Churia Hills, are a mountain range of the outer Himalayas. The literal translation of "Sivalik" is 'tresses of Shiva'. The hills are known for their numerous fossils, and are also home to the Soanian Middle Pale ...
of the northern Indian subcontinent. 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.


Description

The Asian species of ''Hexaprotodon'', like the living
hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...
(''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 In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
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 In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral ha ...
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.
Dental microwear Dental microwear analysis is a method to infer diet and behavior in extinct animals, especially in fossil specimens. It has been used on a variety of taxa, including hominids, victoriapithecids, amphicyonids, canids, ursids, hyaenids, hyaenodont ...
suggests a grazing diet for Asian ''Hexaprotodon'' species, similar to ''H. amphibius''.


Extinction

''Hexaprotodon'' was largely extinct by the Late
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
in Southeast Asia but survived in
Sumatra Sumatra () is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the list of islands by area, sixth-largest island in the world at 482,286.55 km2 (182,812 mi. ...
into the early Late Pleistocene, with one tooth dated to around 70,000 years ago. The last known populations survived on the
Indian Subcontinent The Indian subcontinent is a physiographic region of Asia below the Himalayas which projects into the Indian Ocean between the Bay of Bengal to the east and the Arabian Sea to the west. It is now divided between Bangladesh, India, and Pakista ...
to the very end of the Pleistocene, with among the latest dates being around 16,467–15,660 cal years
Before Present Before Present (BP) or "years before present (YBP)" is a time scale used mainly in archaeology, geology, and other scientific disciplines to specify when events occurred relative to the origin of practical radiocarbon dating in the 1950s. Because ...
from bones found in the
Narmada River The Narmada River, previously also known as ''Narbada'' or anglicised as ''Nerbudda'', is the 5th longest river in India and overall the longest west-flowing river in the country. It is also the largest flowing river in the state of Madhya Prade ...
valley in central India. Fossil evidence from a late-surviving Indian ''Hexaprotodon'' indicates that it lived during a catastrophic
drought A drought is a period of drier-than-normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, ...
caused by the latest
Heinrich event A Heinrich event is a natural phenomenon in which large groups of icebergs break off from the Laurentide ice sheet and traverse the Hudson Strait into the North Atlantic. First described by the marine geologist Hartmut Heinrich, they occurred dur ...
, leading to an extremely weak
Indian monsoon The Monsoon of South Asia is among several geographically distributed global monsoons. It affects the Indian subcontinent, where it is one of the oldest and most anticipated weather phenomena and an economically important pattern every year fro ...
. It is thought that these drought conditions led to a heavy
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological proces ...
due to ''Hexaprotodon'' depending on aquatic habitats, prompting an
extinction vortex Extinction vortices are a class of models through which conservation biologists, geneticists and ecologists can understand the dynamics of and categorize extinctions in the context of their causes. This model shows the events that ultimately lea ...
. 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.


Species

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


References

Extinct hippopotamuses Prehistoric Artiodactyla genera Extinct mammals of Asia Pleistocene Artiodactyla Fossil taxa described in 1836 {{Paleo-eventoedungulate-stub