Equus (genus)

Equus (/ˈɛkwəs, ˈiːkwəs/) is a genus of mammals in the perissodactyl family Equidae, which includes horses, asses, and zebras. Within the Equidae, Equus is the only recognized extant genus, comprising seven living species. Like Equidae more broadly, Equus has numerous extinct species known only from fossils. The genus originated in North America and dispersed into the Old World and South America during the Early and Middle Pleistocene. Equines are odd-toed ungulates with slender legs, long heads, relatively long necks, manes (erect in most subspecies), and long tails. All species are herbivorous, and mostly grazers, with simpler digestive systems than ruminants but able to subsist on lower-quality vegetation.

While the domestic horse and donkey (along with their feral descendants) exist worldwide, wild equine populations are limited to Africa and Asia. Wild equine social systems are in two forms; a harem system with tight-knit groups consisting of one adult male or stallion/jackass, several females or mares/jennets, and their young or foals; and a territorial system where males establish territories with resources that attract females, which associate very fluidly. In both systems, females take care of their offspring, but males may play a role as well. Equines communicate with each other both visually and vocally. Human activities have threatened wild equine populations.

The word equus is Latin for "horse" and is cognate with the Greek ἵππος (hippos, "horse") and Mycenaean Greek i-qo /ikkʷos/, the earliest attested variant of the Greek word, written in Linear B syllabic script. Compare the alternative development of the Proto-Greek labiovelar in Ionic ἴκκος (ikkos).

The genus Equus was first described by Carl Linnaeus in 1758. It is the only recognized extant genus in the family Equidae. The first equids were small, dog-sized mammals (e.g. Eohippus) adapted for browsing on shrubs during the Eocene, around 54 million years ago (Mya). These animals had three toes on the hind feet and four on the front feet with small hooves in place of claws, but also had soft pads. Equids developed into larger, three-toed animals (e.g. Mesohippus) during the Oligocene and Miocene. From there, the side toes became progressively smaller through the Pleistocene until the emergence of the single-toed Equus.

The genus Equus, which includes all extant equines, is believed to have evolved from Dinohippus, via the intermediate form Plesippus. One of the oldest species is Equus simplicidens, described as zebra-like with a donkey-like head shape. The oldest material to date was found in Idaho, USA. The genus appears to have spread quickly into the Old World, with the similarly aged E. livenzovensis documented from western Europe and Russia. Molecular phylogenies indicate that the most recent common ancestor of all modern equines (members of the genus Equus) lived ~5.6 (3.9-7.8) Mya. Direct paleogenomic sequencing of a 700,000-year-old middle Pleistocene horse metapodial bone from Canada implies a more recent 4.07 Mya for the most recent common ancestor within the range of 4.0 to 4.5 Mya.

Mitochondrial evidence supports the division of Equus species into noncaballoid (which includes zebras and asses) and caballoids or "true horses" (which includes E. caballus and E. ferus przewalskii, alternatively E. przewalskii). Of the extant equine species, the lineage of the asses may have diverged first,^{[contradictory]} possibly as soon as Equus reached the Old World. Zebras appear to be monophyletic and differentiated in Africa, where they are endemic. Members of the subgenus Sussemionus were abundant during the Early and Middle Pleistocene of North America and Afro-Eurasia, but only a single species, Equus ovodovi survived into the Late Pleistocene and Holocene in south Siberia and China, with the youngest remains from China dating to around 3500 BP (1500 BC), during the Shang dynasty. Genetic data from E. ovodovi has placed the Sussemionus lineage as closer to zebras and asses than to caballine horses.

Molecular dating indicates the caballoid lineage diverged from the noncaballoids 4 Mya. Genetic results suggest that all North American fossils of caballine equines, as well as South American fossils traditionally placed in the subgenus E. (Amerhippus), belong to E. ferus. Remains attributed to a variety of species and lumped together as New World stilt-legged horses (including E. francisci, E. tau, and E. quinni) probably all belong to a second species that was endemic to North America. This was confirmed in a genetic study done in 2017, which subsumed all the specimens into the species E. francisci which was placed outside all extant horse species in the new genus Haringtonhippus, although its placement as a separate genus was subsequently questioned. A separate genus of horse, Hippidion existed in South America. The possible causes of the extinction of horses in the Americas (about 12,000 years ago) have been a matter of debate. Hypotheses include climatic change and overexploitation by newly arrived humans. Horses only returned to the American mainland with the arrival of the conquistadores in 1519.

Many extinct prehistoric species of Equus have been described. The validity of some of these species is questionable and a matter of debate. For example, Equus niobrarensis is likely synonymous with Equus scotti, while Equus alaskae is most likely the same species as Equus lambei, which itself may be a North American form of the living Equus przewalskii.