
Perissodactyla, as we know it today, is a small
order of hoofed mammals, containing 17 Recent species in three families:
Equidae (horses), Tapiridae (tapirs), and Rhinocerotidae (rhinoceroses).
Despite being such a small order, the odd-toed ungulates have a wide
distribution, being found across the Ethiopian, Palearctic, Oriental, and
Neotropical zoogeographic regions. Two species have been domesticated - the
horse (
Equus caballus) and the donkey (
Equus asinus); feral
populations of these two species have been established across the globe,
(re)expanding this order's influence to the Nearctic and Australasian regions.
All modern perissodactyls are large to very large animals, ranging in size
from the smallest equids (200 kg) to the largest rhinoceroses (3,500 kg).
Despite the broad range in their physical appearances, perissodactyls are
united by their mesaxonic limb structure, with most of the body weight being
borne by the large central digit. Although a small order now, there are fourteen
known families (living and extinct), which can be split into three suborders:
-
Hippomorpha - horse-like perissodactyls.
-
Ceratomorpha - rhinoceroses and relatives, including tapirs.
-
Ancylopoda - the now-extinct chalicotheres.
Pe·ris·so·dac·ty·la
(pai'ris-oh dak ti'lah)
From Greek perissos, strange, of numbers odd; daktulos, a finger
or toe |
Evolution
Perissodactyls evolved on the Mesozoic continent of Laurasia, diversifying
rapidly in what is now North America. The ungulate fauna of the Eocene was
dominated by Perissodactyls, with thirteen different families evolving and
spreading across the globe. Their ranks included some of the largest land
mammals to have ever existed, including the ceratomorph Indricotherium
which stood over 5 meters tall at the shoulder and weighed an estimated 15-20
tons! The diversity of perissodactyls has continually declined since the
Oligocene, simultaneous with the rise in the artiodactyls, or even-toed
ungulates. Only four perissodactyl families (the three extant families, plus
the Chalicotheriidae) survived to the Pleistocene, and only 16 species in
6 genera have survived to the present day (the nominate form of Equus
quagga, the South African quagga, became extinct in 1883 due to hunting and habitat loss, but it is now believed to be synonymous with the plains zebra). Modern
perissodactyls are a last remnant of a once exceptionally successful order,
and today, with the exception of domestic horses and donkeys, all species
are found in relatively low numbers.
Diagnostic Characteristics
In all species, digit III is the most prominent on all feet, and, as the
plane of symmetry of the foot passes through this digit, perissodactyls are
said to have a mesaxonic foot. The first digit (equivalent the thumb or big
toe of humans) is lost in all species. The Equidae have a single functional
toe on each foot (the third digit), while the Rhinocerotidae have three toes
per foot. The Tapiridae - the closest family to the ancestral perissodactyl
condition - possess four toes on the forefeet (digit V is used on wet or
marshy ground) and three on the hind. Perissodactyls are truly unguligrade,
with the heel, sole, and digits of the foot never touching the ground. The
ulna and fibula (bones in the forearm and lower leg) are reduced, simplifying
the wrist/ankle joint considerably. The clavicle (collar bone) is absent,
allowing for efficient running - the main driving force behind unguligrade
evolution.
The skull is elongated, with the expansion occuring as a result of the facial
bones being stretched (rather than the braincase). The nasal bones in the
skull are expanded posteriorly, and project freely for at least part of their
length. There is a well-developed paroccipital process. 'Horns' are present
on the midline of the nasal and/or frontal bones in all living members of
the Rhinocerotidae, but these are dermal in origin and have no bony core
as in the artiodactyls. The dental formula is quite variable among modern
species, I 0-3/0-3, C 0-1/0-1, P 3-4/3-4, M 3/3 x 2 = 24-44. In grazing species
such as the horses the molars and premolars are hypsodont (high-crowned to
allow for wear), while browsing forms such as the tapirs have brachydont
(low-crowned) teeth. The pattern of dental ridges in modern perissodactyls
is lophodont.
The Perissodactyla Family Tree
Branch lengths are not proportional to time
(From Norman and Ashley, 2000)