Hippopotamidae

This is the article on the family Hippopotamidae; for the main article on hippos, see Hippopotamus

Hippopotamidae Temporal range: 16–0 Ma PreЄ Є O S D C P T J K Pg N Early Miocene-Holocene
Common hippopotamus
Pygmy hippopotamus
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Artiodactyla
(unranked)	Whippomorpha
Family:	Hippopotamidae Gray, 1821
Subtaxa
†Trilobophorous Geze, 1985 Hippopotaminae Gray, 1821 †Archaeopotamus Choeropsis †Hexaprotodon Hippopotamus †Saotherium †Kenyapotaminae Pickford, 1983 Kenyapotamus Palaeopotamus

Hippopotamuses are stout, naked-skinned, and amphibious artiodactyl members (and the only living members) of the family Hippopotamidae possessing three-chambered stomachs and walking on four toes on each foot. While they resemble pigs physiologically and are pseudoruminants like camels, their closest living relatives are actually the cetaceans (whales and dolphins).

There are two living species of hippopotamus in two genera; the pygmy hippo, Choeropsis liberiensis of the forests of west Africa, and the common hippo, Hippopotamus amphibius.

Family Hippopotamidae (hippos: 2 genera, 2 species)

•Africa south of Sahara; pygmy hippo only in western Africa

•Hippopotamus up to 3600 kg, Hexaprotodon only about 180 kg

•amphibious, restricted to areas near water but comes on shore at night to graze

•skin glands exude reddish, oily substance; mostly hairless except bristles on snout

•bunodont molars, tusk-like canines

•common hippo gregarious, pygmy hippo solitary or in pairs

Characteristics

Hippopotamuses 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 their snouts. 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 on each foot. Their stomachs have 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 hippos dehydrate rapidly in dry environments.

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

Dentition
2–3.1.4.3
1–3.1.4.3

Evolution

Main article: Evolution of hippopotamus

The hippopotamids are descended from the anthracotheres, a family of semiaquatic and terrestrial artiodactyls that appeared in the late Eocene, and are 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 relationship to each other - apart from being fairly distant relatives - is 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, has turned out to be paraphyletic.

Taxonomy

Artiodactyla	Tylopoda    Artiofabula     Suina        Cetruminantia     Ruminantia    Whippomorpha     Hippopotamidae  Cetacea

Analogous structures

The lower canine teeth of hippopotamids are similar in function and structure to the tusks of elephants. While hippopotamids and elephants are only very distantly related within the Mammalia, the lower canine teeth of both groups are long and have a slight curve, and species of both families 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 and mainland Europe; Late Miocene–Middle Pleistocene
• Tentatively placed into Hippopotamus:
  • †Hippopotamus aethiopicus - Kenya & Ethiopia; Pliocene -Pleistocene
  • †Hippopotamus afarensis - formerly genus Trilobophorus
  • †Hippopotamus behemoth - Israel, Early Pleistocene
  • †Hippopotamus kaisensis - Central Africa; Pliocene
  • †Hippopotamus sirensis - Morocco & Algeria; Pleistocene
• Genus Hexaprotodon - hexaprotodons or Asian hippopotamuses
  • †Hexaprotodon bruneti - Ethiopia; Pliocene
  • †Hexaprotodon coryndoni - Ethiopia; Pliocene
  • †Hexaprotodon crusafonti - Spain; Late Miocene (syn. Hexaprotodon primaevus)
  • †Hexaprotodon hipponensis - Algeria
  • †Hexaprotodon imagunculus - Uganda and Congo; Pliocene
  • †Hexaprotodon iravaticus - Myanmar; Pliocene - Pleistocene
  • †Hexaprotodon karumensis - Kenya and Eritrea; Pleistocene
  • †Hexaprotodon madagascariensis - Madagascar; Holocene
  • †Hexaprotodon namadicus - India; (possibly same as Hex. palaeindicus)
  • †Hexaprotodon palaeindicus - India;
  • †Hexaprotodon pantanellii - Italy; Pliocene
  • †Hexaprotodon protamphibius - Kenya and Chad; Pliocene
  • †Hexaprotodon siculus -
  • †Hexaprotodon sivajavanicus - Indonesia
  • †Hexaprotodon sivalensis - India
  • †Hexaprotodon sp. (undescribed) - Myanmar

• Genus Archaeopotamus - formerly included in Hexaprotodon
  • †Archaeopotamus harvardi - Arabian Peninsula and Central Africa; Miocene - Pliocene
  • †Archaeopotamus lothagamensis - Kenya; Miocene
• One or two undescribed species
• Genus Choeropsis
  • Choeropsis liberiensis - pygmy hippopotamus
• Genus Saotherium - formerly included in Hexaprotodon
  • Saotherium mingoz Chad; Pliocene

Further reading

• 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. doi:10.1111/j.1096-3642.2004.00138.x (HTML abstract)
• Petronio, C. (1995): Note on the taxonomy of Pleistocene hippopotamuses. Ibex 3: 53-55. PDF fulltext