Anthracotheriidae Temporal range: | |
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Anthracotherium | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Artiodactyla |
Suborder: | Whippomorpha |
Superfamily: | Hippopotamoidea |
Family: | † Anthracotheriidae Leidy, 1869 |
Genera and subfamilies [1] [2] | |
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. [3] The youngest genus, Merycopotamus , died out in Asia during the late Pliocene, possibly for the same reasons. 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. [4]
In life, the average anthracothere would have resembled a skinny hippopotamus with a comparatively small, narrow head and most likely pig-like in general appearance. [5] They had four or five toes on each foot, and broad feet suited to walking on soft mud. They had full sets of about 44 teeth with five semicrescentric cusps on the upper molars, [3] which, in some species, were adapted for digging up the roots of aquatic plants. [6]
Some skeletal characters of anthracotheres suggest they are related to hippos. [7] The nature of the sediments in which they are fossilized implies they were amphibious, which supports the view, based on anatomical evidence, that they were ancestors of the hippopotamuses. [8] In many respects, especially the anatomy of the lower jaw, Anthracotherium, as with other members of the family, is allied to the hippopotamus, of which it is probably an ancestral form. [9] However, one study suggests that instead of anthracotheres, another pig-like group of artiodactyls, the palaeochoerids, are the true stem group of Hippopotamidae. [10]
Recent evidence, gained from comparative gene sequencing, further suggests that hippos are the closest living relatives of whales, [11] [12] so, if anthracotheres are stem hippos, they would also be related to whales in a clade provisionally called Whippomorpha.
However, the earliest known anthracotheres appear in the fossil record in the middle Eocene, well after the archaeocetes had already taken up totally aquatic lifestyles. Although phylogenetic analyses of molecular data on extant animals strongly support the notion that hippopotamids are the closest relatives of cetaceans (whales, dolphins and porpoises), the two groups are unlikely to be closely related when extant and extinct artiodactyls are analyzed. Cetaceans originated about 50 million years ago in the Tethys Sea between India and China, whereas the family Hippopotamidae is only 15 million years old, and the first Asian hippopotamids are only 6 million years old. Yet, analyses of fossil clades have not resolved the issue of cetacean relations. [13]
Another study has offered a suggestion that anthracotheres are part of a clade that also consists of entelodonts (and even Andrewsarchus ) and that is a sister clade to other cetancodonts, with Siamotherium as the most basal member of the clade Cetacodontamorpha. [14]
Ungulates are members of the diverse clade Euungulata which primarily consists of large mammals with hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined to be a polyphyletic and thereby invalid clade based on molecular data. As a result, true ungulates had since been reclassified to the newer clade Euungulata in 2001 within the clade Laurasiatheria while Paenungulata has been reclassified to a distant clade Afrotheria. Living ungulates are divided into two orders: Perissodactyla including equines, rhinoceroses, and tapirs; and Artiodactyla including cattle, antelope, pigs, giraffes, camels, sheep, deer, and hippopotamuses, among others. Cetaceans such as whales, dolphins, and porpoises are also classified as artiodactyls, 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, both native to South America, became extinct at the end of the Pleistocene, around 12,000 years ago.
The hippopotamus, also shortened to 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).
Artiodactyls are placental mammals belonging to the order Artiodactyla. Typically, they are ungulates which bear weight equally on two of their five toes: the third and fourth, often in the form of a hoof. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of the five toes. Another difference between the two is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine as perissodactyls do. The advent of molecular biology, along with new fossil discoveries, found that cetaceans fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use "even-toed ungulates" to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature.
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 sometimes referred to as hippopotamids.
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.
The evolution of cetaceans is thought to have begun in the Indian subcontinent from even-toed ungulates (Artiodactyla) 50 million years ago (mya) and to have proceeded over a period of at least 15 million years. Cetaceans are fully aquatic marine mammals belonging to the order Artiodactyla and branched off from other artiodactyls around 50 mya. Cetaceans are thought to have evolved during the Eocene, the second epoch of the present-extending Cenozoic Era. Molecular and morphological analyses suggest Cetacea share a relatively recent closest common ancestor with hippopotami and that they are sister groups. Being mammals, they surface to breathe air; they have 5 finger bones (even-toed) in their fins; they nurse their young; and, despite their fully aquatic life style, they retain many skeletal features from their terrestrial ancestors. Research conducted in the late 1970s in Pakistan revealed several stages in the transition of cetaceans from land to sea.
Mesonychia is an extinct taxon of small- to large-sized carnivorous ungulates related to artiodactyls. Mesonychians first appeared in the early Paleocene, went into a sharp decline at the end of the Eocene, and died out entirely when the last genus, Mongolestes, became extinct in the early Oligocene. In Asia, the record of their history suggests they grew gradually larger and more predatory over time, then shifted to scavenging and bone-crushing lifestyles before the group became extinct.
Anthracotherium is an extinct genus of artiodactyls characterized by having 44 teeth, with five semi-crescentic cusps on the crowns of the upper molars. The genus ranged from the middle Eocene period until the early Miocene, having a distribution throughout Eurasia. Material subjectively assigned to Anthracotherium from Pakistan suggests the last species died out soon after the start of the Miocene.
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 artiodactyls that contains all living cetaceans and hippopotamuses. 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.
Sinonyx is a genus of extinct, superficially wolf-like mesonychid mammals from the late Paleocene of China. It is within the family Mesonychidae, and cladistic analysis of a skull of Sinonyxjiashanensis identifies its closest relative as Ankalagon. S.jiashanensis was discovered in Anhui province, China, in the Tuijinshan formation.
Indohyus is an extinct genus of digitigrade even-toed ungulates known from Eocene fossils in Asia. This small chevrotain-like animal found in the Himalayas is one of the earliest known non-cetacean ancestors of whales.
Anthracothema was a genus of extinct artiodactyl ungulate mammals that lived in Myanmar during the late Eocene.
Artiocetus is an extinct genus of early whales belonging to the family Protocetidae. It was a close relative to Rodhocetus and its tarsals indicate it resembled an artiodactyl.
Artiofabula is a clade made up of the Suina and the Cetruminantia. The clade was found in molecular phylogenetic analyses and contradicted traditional relationships based on morphological analyses.
Microbunodon was a genus of extinct artiodactyl mammals in the family Anthracotheriidae. It lived between the upper Eocene and the lower Pliocene. Its fossil remains have been found in Europe and Asia.
The bothriodontines are a paraphyletic assemblage of anthracotheres that originated from Eurasia in the late middle Eocene (Bartonian). The group can be distinguished from other anthracothere lineages by their upper molars with the mesostyle that is occupied by the transverse valley, selenodont cusps, ventrally concave symphysis, elongated muzzles, with presence of a diastema between the canine and first premolar. The size range of the group ranged from small, basal forms to larger and more derived forms. During their evolution, the bothriodontines undergone a trend from evolving from small basal forms such as Qatraniodon into larger taxa such as Libycosaurus and Merycopotamus. Some genera the snouts became even more elongated and teeth specialized in a folivorous diet, while others like Merycopotamus became wide, heavy and shallow muzzles with teeth more adapted for grazing.
The microbunodontines were an extinct subfamily of anthracotheres that were predominately a Paleogene group of Eurasian artiodactyls. The group died out at the end of the Late Miocene. It comprised the genera Anthracokeryx, Geniokeryx, Microbunodon, and possibly Etruscotherium. They are different from the other anthracothere lineages by their smaller size, slenderer limbs and male specimens having laterally compressed, longer canines. They were originally classified as members of the other subfamily of anthracotheres, Anthracotheriinae but recent phylogenetic studies have found them to be their own clade sister to Bothriodontinae.
The anthracotheriines are an extinct subfamily of anthracotheres that comprised Paleogene to early Neogene North American and Eurasian artiodactyls. The group contained the genera Anthracotherium, Heptacodon, and Paenanthracotherium, as well as possibly Myaingtherium and Siamotherium. They were small to large sized anthracotheres, and when compared to the other two subfamilies, Microbunodontinae and Bothriodontinae, anthracotheriines are found to occupy a primitive, basal position in the family.
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.