Hippopotamidae Temporal range: Miocene-Recent Late | |
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Common hippopotamus | |
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Pygmy hippopotamus | |
Scientific classification ![]() | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Suborder: | Whippomorpha |
Superfamily: | Hippopotamoidea |
Family: | Hippopotamidae Gray, 1821 |
Subtaxa | |
† Trilobophorus Geze, 1985
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Hippopotamidae is a family of stout, naked-skinned, and semiaquatic artiodactyl mammals, possessing three-chambered stomachs and walking on four toes on each foot. While they resemble pigs physiologically, their closest living relatives are the cetaceans. They are formally referred to as hippopotamids.
There are two living species of hippopotamid in two genera; the pygmy hippo, Choeropsis liberiensis of the forests of west Africa, and the common hippo, Hippopotamus amphibius . The term hippopotamus can also be applied to hippopotamids in general, although it is most frequently used for the common hippo and its respective genus.
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 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. [1]
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 given in the table below:
Dentition |
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2–3.1.4.3 |
1–3.1.4.3 |
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. The hippos split off from the anthracotheres some time during the Miocene. The oldest records of Hippopotamidae are from Afro-Arabia and date to the late Miocene, approximately 7.4 million years ago, expanding into Eurasia around 6 million years ago. [2] After the appearance of the hippopotamids, the remaining 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.
Hippopotamidae's placement within Artiodactyla can be represented in the following cladogram: [3] [4] [5] [6] [7]
Artiodactyla |
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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.
The systematics and taxonomy used here mostly follows a review by J.-R. Boisserie [8] [lower-alpha 1] and the American Society of Mammalogists. [9]
Ungulates are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. Living ungulates are divided into two orders: the odd-toed ungulates (Perissodactyla) including horses, rhinoceroses, and tapirs; and even-toed ungulates (Artiodactyla) such as cattle, pigs, giraffes, camels, sheep, deer, and hippopotamuses. Cetaceans such as whales, dolphins, and porpoises are also classified as even-toed ungulates, although they do not have hooves. Most terrestrial ungulates use the hoofed tips of their toes to support their body weight while standing or moving. Two other orders of ungulates, Notoungulata, and Litopterna, native to South America, became extinct at the end of the Pleistocene, around 12,000 years ago.
The hippopotamus or hippo, further qualified as the common hippopotamus, Nile hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extant species in the family Hippopotamidae, the other being the pygmy hippopotamus. Its name comes from the ancient Greek for "river horse" (ἱπποπόταμος).
The even-toed ungulates are ungulates—hoofed animals—which bear weight equally on two of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, odd-toed ungulates bear weight on an odd number of the five toes. Another difference between the two is that many other even-toed ungulates digest plant cellulose in one or more stomach chambers rather than in their intestine as the odd-toed ungulates do.
The pygmy hippopotamus or pygmy hippo is a small hippopotamid which is native to the forests and swamps of West Africa, primarily in Liberia, with small populations in Sierra Leone, Guinea, and Ivory Coast. It has been extirpated from Nigeria.
Crocodylus is a genus of true crocodiles in the family Crocodylidae.
Hippopotamus is a genus of artiodactyl mammals consisting of one extant species, Hippopotamus amphibius, the river hippopotamus, and several extinct species from both recent and prehistoric times. It belongs to the family Hippopotamidae, which also includes the pygmy hippopotamus and a number of extinct genera.
Hexaprotodon is an extinct genus of hippopotamid known from Africa and Asia. The name Hexaprotodon means "six front teeth" as some of the fossil forms have three pairs of incisors.
Anthracotheriidae is a paraphyletic family of extinct, hippopotamus-like artiodactyl ungulates related to hippopotamuses and whales. The oldest genus, Elomeryx, first appeared during the middle Eocene in Asia. They thrived in Africa and Eurasia, with a few species ultimately entering North America during the Oligocene. They died out in Europe and Africa during the Miocene, possibly due to a combination of climatic changes and competition with other artiodactyls, including pigs and true hippopotamuses. The youngest genus, Merycopotamus, died out in Asia during the late Pliocene. The family is named after the first genus discovered, Anthracotherium, which means "coal beast", as the first fossils of it were found in Paleogene-aged coal beds in France. Fossil remains of the anthracothere genus were discovered by the Harvard University and Geological Survey of Pakistan joint research project (Y-GSP) in the well-dated middle and late Miocene deposits of the Pothohar Plateau in northern Pakistan.
Kenyapotamus is a possible ancestor of living hippopotamuses that lived roughly 16 million to 8 million years ago during the Miocene epoch. Its name reflects that its fossils were first found in modern-day Kenya.
Whippomorpha or Cetancodonta is a group of animals that contains all living cetaceans and hippopotamuses, as well as their extinct relatives, i.e. Entelodonts and Andrewsarchus. All Whippomorphs are descendants of the last common ancestor of Hippopotamus amphibius and Tursiops truncatus. This makes it a crown group. Whippomorpha is a suborder within the order Artiodactyla. The placement of Whippomorpha within Artiodactyla is a matter of some contention, as hippopotamuses were previously considered to be more closely related to Suidae (pigs) and Tayassuidae (peccaries). Most contemporary scientific phylogenetic and morphological research studies link hippopotamuses with cetaceans, and genetic evidence has overwhelmingly supported an evolutionary relationship between Hippopotamidae and Cetacea. Modern Whippomorphs all share a number of behavioural and physiological traits; such as a dense layer of subcutaneous fat and largely hairless bodies. They exhibit amphibious and aquatic behaviors and possess similar auditory structures.
The Cetruminantia are a clade made up of the Cetancodontamorpha and their closest living relatives, the Ruminantia.
The Cyprus dwarf hippopotamus or Cypriot pygmy hippopotamus is an extinct species of hippopotamus that inhabited the island of Cyprus from the Pleistocene until the early Holocene.
Hippopotamus pentlandi is an extinct species of hippopotamus from Sicily, known from the late Middle Pleistocene to early Late Pleistocene. It is the largest of the insular dwarf hippos known from the Pleistocene of the Mediterranean, "at most 20% smaller than the mainland forms", with an estimated body mass of approximately 1100 kg. It is suggested that it arrived in Sicily between 250,000 and 150,000 years ago, probably descending from the modern hippopotamus, with an origin from Hippopotamus antiquus being less likely. In comparison to those species, the muzzle was shorter, the occipital and nasal regions were more developed, the mastoid process was enlarged, and the dental row was shortened, and the condyle of the mandible is low. In comparison to H. amphibius, the orbits are also elevated. It was present in Sicily until at least the latest Middle Pleistocene around 120 kya, and was probably extinct by the beginning of Marine Isotope Stage 4. Contemporaneous species include the dwarf elephant Palaeoloxodon mnaidriensis. Its diet was likely grazing dominated, similar to that of modern H. amphibius. It is probably ancestral to Hippopotamus melitensis from Malta, which is substantially smaller than H. pentlandi.
Lemerle's dwarf hippopotamus is an extinct species of Malagasy hippopotamus.
Archaeopotamus is an extinct genus of Hippopotamidae that lived between 7.5 and 2.58 million years ago in Africa and the Middle East. The genus was described in 2005 to encompass species of hippos that were previously grouped in Hexaprotodon.
Several species of Malagasy hippopotamus lived on the island of Madagascar but are now believed to be extinct. The animals were very similar to the extant hippopotamus and pygmy hippopotamus. The fossil record suggests that at least one species of hippopotamus lived until about 1,000 years ago and other evidence suggests that the species may have survived until much more recently. The taxonomy of these animals is not resolved and not widely studied. The various species are believed to have survived into the Holocene epoch.
Hippopotamus laloumena is an extinct species of hippopotamus from Pleistocene and Holocene Madagascar, making it the oldest of Malagasy hippopotamus. H. laloumena was much larger than other Malagasy hippopotamus, but was still somewhat smaller than the common hippopotamus. However, little is known about the species because it was identified with only a lower jaw and limb bones. It was described in 1990 by French palaeontologists M. Faure and Guerin, the fossils recovered from a site near Mananjary on the east coast of Madagascar. The species name derives from Malagasy laloumena "hippopotamus".
Hippopotamus madagascariensis, the Madagascar or Madagascan dwarf hippopotamus, is an extinct species of hippopotamus, endemic to the island of Madagascar. Known only from bones, it is believed to have gone extinct sometime after 1500. It was one of three hippopotamus species, the Malagasy hippopotamuses, which were native to Madagascar and are all extinct.
Saotherium is an extinct genus of hippopotamid from the Early Pliocene of Africa, specifically Chad. It is represented by a single species, S. mingoz. The earliest fossils appear at the start of the Pliocene, while the latest are dated to about 4 mya.
Ancodonta is an infraorder of semiaquatic artiodactyl ungulates including modern hippopotamus and all mammals closer to hippos than to cetaceans (whales). Ancodonts first appeared in the Middle Eocene, with some of the earliest representatives found in fossil deposits in Southeast Asia. Throughout their evolutionary history they have occupied different browsing and grazing niches in North America, Eurasia and Africa. The last continent is notable as they were among the first laurasiatherian mammals to have migrated to Africa from Europe, where they competed with the native afrothere herbivores for the same niches. Of the nearly 50 genera that have existed, only two of them are extant – Choeropsis and Hippopotamus. The interrelationships within the ancodonts has been contended. The traditional notion is that there at minimum two families Anthracotheriidae and Hippopotamidae and were merely sister taxa. However many detailed research of the dentition among ancodonts, as well as how some anthracotheres were similar to hippos in appearance, lead the current consensus where Anthracotheriidae is paraphyletic to Hippopotamidae. Among the anthracotheres members of Bothriodontinae are among the closest to the ancestry of hippos, with the Oligocene aged Epirigenys from Lokon, Turkana, Kenya being the sister taxon to hippos. In response of this many similar clade names have been used for this clade.