Evolution and paleontology

The artiodactyls can be traced back to a probable descent from a group of early generalized mammals called condylarths and were certainly distinct by the Eocene Epoch, which ended about 33.9 million years ago. Fossil artiodactyls can be more or less convincingly classified in three suborders; the more primitive Suiformes, centred around pigs, the Tylopoda, centred on camels, and the Ruminantia or ruminants. The most primitive artiodactyls are the suiform group Palaeodonta, which had four functional toes on each foot, primitive, low-cusped cheek teeth, and the typical artiodactyl astragalus. The artiodactyls became more prominent in the Oligocene (between about 33.9 million and 23 million years ago) with a decline of the then dominant perissodactyls, and the later history of artiodactyls appears as successive waves of groups, each better adapted than its predecessors to the changing environment. In the suiform line, the earlier palaeodonts are succeeded by other groups such as the entelodonts, giant “pigs” of the European and North American Oligocene, characterized by very large skulls (some nearly 1 metre [3.3 feet] long), very small brains, and a large, bony flange below the eyes. The functionally two-toed ruminants succeeded four-toed suiforms in the Miocene, and within the Old World ruminants of the bovid subfamily Caprinae, the zenith of the tribe Caprini, for example, followed that of the mainly Pliocene tribe Ovibovini.

The artiodactyls had an interesting history in North America through the Paleogene and Neogene periods. Some forms, such as the entelodonts, were shared with the Old World, but others were characteristic of North America. One very prominent New World family was the merycoidodonts (or oreodonts), which lasted until the early Pliocene (about 3.6 million years ago). They had somewhat piglike proportions, short faces, a large upper canine and a caniniform first lower premolar, and selenodont molars. A close relative, Agriochoerus, had clawed feet, the function of which remains uncertain.

Camelids evolved in North America and, at or toward the end of the Neogene, spread into South America and into the Old World. By the end of the Pleistocene they all became extinct in their homeland, just as horses did. The hypertragulids were a mainly Oligocene group of chevrotain-like forms related to the Protoceratidae. The latter had horns above their noses, a position unique among artiodactyls, as well as in the usual position. The North American Miocene (23 million to 5.3 million years ago) produced some ruminants, such as Blastomeryx, that are hard to distinguish from the early palaeomerycine relatives of giraffes and deer in the Old World, which, with the North American groups, constitute the family Palaeomerycidae. Some developed horns, and the dromomerycine Cranioceras even had a third horn above the back of its skull. During the Miocene and Pliocene there finally appeared relatives of the surviving pronghorn, an example being Merycodus. Many of these North American groups have parallels with Old World groups, and the subject of North American artiodactyl evolution is of great interest. Only further finds will indicate whether Blastomeryx, the dromomerycines, Merycodus, and the pronghorns evolved from hypertragulids already in North America or sprang from some immigrant ruminant and, if the latter, whether the supposed hypertragulid Leptomeryx could be such an immigrant ruminant. It is uncertain whether the hypertragulids are nearer the tragulines or the camels, and how close the oreodonts are to the anthracotheres. Of the great New World radiation there survived after the Pleistocene only three or four camelid species and the pronghorn (deer and bovids in the Americas are immigrants), whereas in the Old World as little as 200 years ago, Eurasia and Africa had abundant deer and antelopes.

Until the Miocene there were some archaic artiodactyls in Europe, the xiphodonts, which have cautiously been taken as tylopods, and the cainotheres and anoplotheres, which are classified near anthracotheres.

A possible ruminant ancestor was Archaeomeryx from the upper Eocene of China, a small animal that already had a fused naviculo-cuboid bone in the ankle. Tragulids occurred in Africa and Eurasia back to the Miocene, and the more advanced gelocids are known from the upper Eocene and lower Oligocene. At the end of the Oligocene, the first ruminants began to appear with teeth more advanced than those of tragulids. From early in the Miocene they began to be recognizable as giraffes, deer, or antelopes, although the last were relatively uncommon before the late Miocene. Much remains to be learned about the detailed early history of these groups. Several different giraffids lived in later Miocene and early Pliocene times, but the group has since declined to only two species. Deer gradually acquired more complicated antlers, which became very large in some lineages. Different subfamilies of bovids originated in Eurasia and Africa, and it is of zoogeographic interest that representatives of African subfamilies have been found as fossils in northern India and Pakistan.

Classification