Hippopotamidae

Hippopotamuses are the members of the family Hippopotamidae. They are the only extant artiodactyls which walk on four toes on each foot.

Characteristics
Hippopotamids are large mammals, with short, stumpy legs, and barrel-shaped bodies. They have large heads, with broad mouths, and nostrils placed at the top of the snout. Like pigs, they have four toes, but unlike pigs, all of the toes are used in walking. Hippopotamids are unguligrade, although, unlike most other such animals, they have no hooves, instead using a pad of tough connective tissue. Their stomach has three chambers, but they are not true ruminants.

The living species are smooth-skinned and lack both sebaceous glands and sweat glands. The outer epidermis is relatively thin, so that hippos dehydrate rapidly in dry environments.

Both the incisors and canines are large and tusk-like, although the canine tusks are by far the largest. The tusks grow throughout life. The postcanine teeth are large and complex, suited for chewing the plant matter that composes their diet. The number of incisors varies even within the same species, but the general dental formula is:

Evolution
The hippopotamids are descended from the anthracotheres, a family of semi-aquatic artiodactyls that appeared in the late Eocene, and are popularly thought to have resembled small- or narrow-headed hippos. More specifically, the hippos split off from the anthracotheres some time during the Miocene. After the appearance of the true hippopotamids, the anthracotheres went into a decline brought about by a combination of climatic change and competition with their descendants, until the last genus, Merycopotamus, died out in the early Pliocene of India.

There were once many species of hippopotamid, but only two survive today: Hippopotamus amphibius, and Choeropsis liberiensis. They are the last survivors of two major evolutionary lineages, the hippos proper and the pygmy hippos, respectively; these lineages could arguably be considered subfamilies but their relationships to each other - apart from being fairly distant relatives - are not well resolved.

The enigmatic Miocene Kenyapotamus is insufficiently known to be assigned a place in the hippo phylogeny with any degree of certainty. In addition, the genus Hexaprotodon, in a sense now restricted to an extinct group of animals once living around the northern and northeastern Indian Ocean, which formerly included most ancient hippos, turned out to be paraphyletic.

Analogous structures
An analogous structure that hippopotamids have is the lower canine teeth, which are similar in function and structure to an elephant’s tusks. Hippopotamids and elephants are not closely related, but the lower canine teeth of the hippopotamus and the tusk of the elephant are both long and have a slight curve and both animals use this structure when fighting.

Species
The systematics and taxonomy used here mostly follows the review of Boisserie (2005).


 * Genus Hippopotamus - true hippopotami
 * Hippopotamus amphibius - Hippopotamus Africa
 * †Hippopotamus antiquus - mainland Europe & British Isles; Pleistocene
 * †Hippopotamus creutzburgi - Crete; Pleistocene
 * †Hippopotamus minor - Cyprus; Pleistocene to Holocene
 * †Hippopotamus melitensis - Malta; Pleistocene
 * †Hippopotamus pentlandi - Sicily; Pleistocene
 * †Hippopotamus lemerlei - Madagascar; Holocene
 * †Hippopotamus laloumena - Madagascar; Holocene
 * †Hippopotamus gorgops - Africa & mainland Europe; Late Miocene–Middle Pleistocene
 * Tentatively placed into Hippopotamus:
 * †Hippopotamus aethiopicus - Kenya & Ethiopia; Pliocene -Pleistocene
 * †Hippopotamus afarensis - formerly genus Trilobophorus
 * †Hippopotamus behemoth -
 * †Hippopotamus kaisensis - Central Africa; Pliocene
 * †Hippopotamus sirensis - Morocco & Algeria; Pleistocene
 * Genus Hexaprotodon - hexaprotodons or Asian hippopotami
 * †Hexaprotodon bruneti - Ethiopia; Pliocene
 * †Hexaprotodon coryndoni - Ethiopia; Pliocene
 * †Hexaprotodon crusafonti - Spain; Late Miocene (syn. Hexaprotodon primaevus)
 * †Hexaprotodon hipponensis - Algeria
 * †Hexaprotodon imagunculus - Uganda & Congo; Pliocene
 * †Hexaprotodon iravaticus - Myanmar; Pliocene - Pleistocene
 * †Hexaprotodon karumensis - Kenya & Eritrea; Pleistocene
 * †Hexaprotodon madagascariensis - Madagascar; Holocene
 * †Hexaprotodon namadicus - India; (possibly same as Hex. palaeindicus)
 * †Hexaprotodon palaeindicus - India;
 * †Hexaprotodon pantanellii - Italy; Pliocene
 * †Hexaprotodon protamphibius - Kenya & Chad; Pliocene
 * †Hexaprotodon siculus -
 * †Hexaprotodon sivalensis - India
 * †Hexaprotodon sp. (undescribed) - Myanmar
 * †Undescribed Indonesian species; Pleistocene


 * Genus Archaeopotamus - formerly included in Hexaprotodon
 * †Archaeopotamus harvardi - Arabian Peninsula& Central Africa; Miocene - Pliocene
 * †Archaeopotamus lothagamensis - Kenya; Miocene
 * 1-2 undescribed species
 * Genus Choeropsis - pygmy hippopotamus
 * Choeropsis liberiensis - Pygmy Hippopotamus
 * Genus Saotherium - formerly included in Hexaprotodon
 * Saotherium mingoz Chad; Pliocene

References and notes

 * Boisserie, Jean-Renaud (2005): The phylogeny and taxonomy of Hippopotamidae (Mammalia: Artiodactyla): a review based on morphology and cladistic analysis. Zool. J. Linn. Soc. 143(1): 1-26. (HTML abstract)
 * Petronio, C. (1995): Note on the taxonomy of Pleistocene hippopotamuses. Ibex 3: 53-55. PDF fulltext
 * Petronio, C. (1995): Note on the taxonomy of Pleistocene hippopotamuses. Ibex 3: 53-55. PDF fulltext

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