Perissodactyl

Odd-toed ungulates, mammals which constitute the taxonomic order Perissodactyla (, ), are animals—ungulates—who have reduced the weight-bearing toes to three (rhinoceroses and tapirs, with tapirs still using four toes on the front legs) or one (equines, third toe) of the five original toes. 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 four or two (an even number) 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 even-toed ungulates, with the exception of Suina, do.

The order includes about 17 species divided into three families: Equidae (horses, asses, and zebras), Rhinocerotidae (rhinoceroses), and Tapiridae (tapirs).

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.

Anatomy

The largest odd-toed ungulates are rhinoceroses, and the extinct '' Paraceratherium'', a hornless rhino from the Oligocene, is considered one of the largest land mammals of all time. At the other extreme, an early member of the order, the prehistoric horse '' Eohippus'', had a withers height of only . Apart from dwarf varieties of the domestic horse and donkey, perissodactyls reach a body length of and a weight of . While rhinos have only sparse hair and exhibit a thick epidermis, tapirs and horses have dense, short coats. Most species are grey or brown, although zebras and young tapirs are striped.

Limbs

The main axes of both the front and rear feet pass through the third toe, which is always the largest. The remaining toes have been reduced in size to varying degrees. Tapirs, which are adapted to walking on soft ground, have four toes on their fore feet and three on their hind feet. Living rhinos have three toes on both the front and hind feet. Modern equines possess only a single toe; however, their feet are equipped with hooves, which almost completely cover the toe. Rhinos and tapirs, by contrast, have hooves covering only the leading edge of the toes, with the bottom being soft.

Ungulates have stances that require them to stand on the tips of their toes. Equine ungulates with only one digit or hoof have decreased mobility in their limb which allows for faster running speeds and agility.
Differences in limb structure and physiology between ungulates and other mammals can be seen in the shape of the humerus. For example, often shorter, thicker, bones belong to the largest and heaviest ungulates like the Rhinoceros.

The ulnae and fibulae are reduced in horses. A common feature that clearly distinguishes this group from other mammals is the articulation between the astragalus, the scaphoid and the cuboid, which greatly restricts the mobility of the foot. The thigh is relatively short, and the clavicle is absent.

Skull and teeth

Odd-toed ungulates have a long upper jaw with an extended diastema between the front and cheek teeth, giving them an elongated head. The various forms of snout between families are due to differences in the form of the premaxilla. The lacrimal bone has projecting cusps in the eye sockets and a wide contact with the nasal bone. The temporomandibular joint is high and the mandible is enlarged.

Rhinos have one or two horns made of agglutinated keratin, unlike the horns of even-toed ungulates, which have a bony core.

The number and form of the teeth vary according to diet. The incisors and canines can be very small or completely absent, as in the two African species of rhinoceros. In the horses, usually only the males possess canines. The surface shape and height of the molars is heavily dependent on whether soft leaves or hard grass make up the main component of their diets. Three or four cheek teeth are present on each jaw half, so the dental formula of odd-toed ungulates is:

Gut

All perissodactyls are hindgut fermenters. In contrast to ruminants, hindgut fermenters store digested food that has left the stomach in an enlarged cecum, where the food is digested by bacteria. No gallbladder is present. The stomach of perissodactyls is simply built, while the cecum accommodates up to in horses. The intestine is very long, reaching up to in horses. Extraction of nutrients from food is relatively inefficient, which probably explains why no odd-toed ungulates are small; nutritional requirements per unit of body weight are lower for large animals, as their surface-area-to-volume ratio is smaller.

Distribution

The present distribution of most perissodactyl species is only a small fraction of their original range. Members of this group are now found only in Central and South America, eastern and southern Africa, and central, southern, and southeastern Asia. During the peak of odd-toed ungulate existence, from the Eocene to the Oligocene, perissodactyls were distributed over much of the globe, the only exceptions being Australia and Antarctica. Horses and tapirs arrived in South America after the formation of the Isthmus of Panama in the Pliocene