In addition to hooves, ungulates developed reduced canine teeth, bunodont molars (molars with low, rounded cusps), and an astragalus (one of the ankle bones at the end of the lower leg) with a short, robust head.
Ungulates diversified rapidly in the Eocene, but are thought to date back perhaps as far as the Late Cretaceous. Most ungulates are herbivores, but a few are omnivores or predators (for example, whales).
Modern ungulates have taken this to the extreme: the metapodials (the bones between the wrist/ankle and the digits) are often as long as the other parts of the legs.
Since the hoof was the defining character of the ungulates, feet were the focus of researchers trying to decipher their origins.
Originally placed at the base of the ungulate lineage, continued research suggested that the paenungulates were more specialized than the true ungulates, nesting them firmly in the ungulate family tree.
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