Order
Perissodactyla
Odd-toed
ungulates |
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:
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Hippomorpha - horse-like perissodactyls.
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Ceratomorpha - rhinoceroses and relatives, including tapirs.
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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 (a 17th species, the quagga Equus quagga became extinct in 1883
due to hunting and habitat loss). 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 Perissodactyl
Family Tree
(branch lengths are not proportional to
time)
Click on each family above to learn
more,
or jump to the Perissodactyla
Species List
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Literature Cited
Klingel, H and E. Thenius. 1990. Odd-toed Ungulates.
In Grzimek's Encyclopedia of Mammals. Volume 4. Edited by
S. P. Parker. McGraw-Hill, New York. Pp. 547-556.
Nowak, R. M. [Editor]. 1991. Walker's Mammals of the World. Fifth Edition.
Johns Hopkins University Press, Baltimore.
Vaughan, T. A., J. M. Ryan, and N. J. Czaplewski. 2000. Mammalogy. Fourth
Edition. Saunders College Publishing, Philadelphia.
Webb, J. E., J. A. Wallwork, and J. H. Elgood. 1979. Guide to Living Mammals.
Second Edition. Bell and Blain Ltd., Glasgow.
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