was called hippotigris ("horse tiger") by the Greeks and Romans. The word "zebra" was traditionally pronounced with a long initial vowel, but over the course of the 20th century the pronunciation with the short initial vowel became the norm in the UK and the Commonwealth. The pronunciation with a long initial vowel remains standard in US English. Taxonomy Zebras are classified in the genus Equus (known as equines) along with horses and asses. These three groups are the only living members of the family Equidae. The plains zebra and mountain zebra were traditionally placed in the subgenus Hippotigris (C. H. Smith, 1841) in contrast to the Grévy's zebra which was considered the sole species of subgenus Dolichohippus (Heller, 1912). Groves and Bell (2004) placed all three species in the subgenus Hippotigris. A 2013 phylogenetic study found that the plains zebra is more closely related to Grévy's zebras than mountain zebras. The extinct quagga was originally classified as a distinct species. Later genetic studies have placed it as the same species as the plains zebra, either a subspecies or just the southernmost population. Molecular evidence supports zebras as a monophyletic lineage. Equus originated in North America and direct paleogenomic sequencing of a 700,000-year-old middle Pleistocene horse metapodial bone from Canada implies a date of 4.07 million years ago (mya) for the most recent common ancestor of the equines within the range of 4.0 to 4.5 mya. Horses split from asses and zebras around this time and equines colonised Eurasia and Africa around 2.1–3.4 mya. Zebras and asses diverged from each other close to 2 mya. The mountain zebra diverged from the other species around 1.6 mya and the plains and Grévy's zebra split 1.4 mya. The cladogram of Equus below is based on Vilstrup and colleagues (2013) and Jónsson and colleagues (2014): Extant species Fossil record In addition to the three living species, some fossil zebras and relatives have also been identified. Equus koobiforensis is an early equine basal to zebras found in the Shungura Formation, Ethiopia and the Olduvai Gorge, Tanzania, and dated to around 2.3 mya. E. oldowayensis is identified from remains in Olduvai Gorge dating to 1.8 mya. Fossil skulls of E. mauritanicus from Algeria which date to around 1 mya appears to show affinities with the plains zebra. E. capensis, known as the Cape zebra, appeared around 2 mya and lived throughout southern and eastern Africa. Non-African