page_title: Odd-toed ungulate
section_title: Research history
hierarchical_section_title: Odd-toed ungulate / Research history
caption_attribution_description: English: Portrait of English anatomist and palaeontologist Richard Owen with the skull of a crocodile. Albumen print from wet collodion on glass negative, 1856.
context_page_description: Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla, are hoofed animals—ungulates—which bear most of their weight on one of the five toes: the third toe. The non-weight-bearing toes are either present, absent, vestigial, or positioned posteriorly. By contrast, the even-toed ungulates bear most of their weight equally on two of the five toes: their third and fourth toes. Another difference between the two is that odd-toed ungulates digest plant cellulose in their intestines rather than in one or more stomach chambers as the even-toed ungulates do.
  The order includes about 17 species divided into three families: Equidae, Rhinocerotidae, and Tapiridae.
  Despite their very different appearances, they were recognized as related families in the 19th century by the zoologist Richard Owen, who also coined the order name.
context_section_description: In 1758, in his seminal work Systema Naturae, Linnaeus (1707–1778) classified horses (Equus) together with hippos (Hippopotamus). At that time, this category also included the tapirs (Tapirus), more precisely the lowland or South American tapir (Tapirus terrestus), the only tapir then known in Europe. Linnaeus classified this tapir as Hippopotamus terrestris and put both genera in the group of the Belluae ("beasts"). He combined the rhinos with the Glires, a group now consisting of the lagomorphs and rodents. Mathurin Jacques Brisson (1723–1806) first separated the tapirs and hippos in 1762 with the introduction of the concept le tapir. He also separated the rhinos from the rodents, but did not combine the three families now known as the odd-toed ungulates. In the transition to the 19th century, the individual perissodactyl genera were associated with various other groups, such as the proboscidean and even-toed ungulates. In 1795, Étienne Geoffroy Saint-Hilaire (1772–1844) and Georges Cuvier (1769–1832) introduced the term "pachyderm" (Pachydermata), including in it not only the rhinos and elephants, but also the hippos, pigs, peccaries, tapirs and hyrax . The horses were still generally regarded as a group separate from other mammals and were often classified under the name Solidungula or Solipèdes, meaning "one-hoof animal".
  In 1861, Henri Marie Ducrotay de Blainville (1777–1850) classified ungulates by the structure of their feet, differentiating those with an even number of toes from those with an odd number. He moved the horses as solidungulate over to the tapirs and rhinos as multungulate animals and referred to all of them together as onguligrades à doigts impairs, coming close to the concept of the odd-toed ungulate as a systematic unit. Richard Owen (1804–1892) quoted Blainville in his study on fossil mammals of the Isle of Wight and introduced the name Perissodactyla.
  In 1884, Othniel Charles Marsh (1831–1899) came up with the concept Mesaxonia, which he used for what are today called the odd-toed ungulates, including their extinct relatives, but explicitly excluding the hyrax. Mesaxonia is now considered a synonym of Perissodactyla, but it was sometimes also used for the true odd-toed ungulates as a subcategory (rhinos, horses, tapirs), while Perissodactyla stood for the entire order, including the hyrax. The assumption that hyraxes were Perissodactyla was held well into the 20th century. Only with the advent of molecular genetic research methods had it been recognized that the hyrax is not closely related to perissodactyls but rather to elephants and manatees.
caption: Richard Owen, 1856
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