The hoofed mammals were present in fairly bewildering variety, but were restricted to the two orders of the Perissodactyla and Artiodactyla. The Perissodactyla, while they no longer had the relatively dominant position which they held in the middle Eocene (see [p. 270]), had suffered no actual loss; and no less than seven families of them, or six by another scheme of classification, had members in the North America of White River times, a very notable difference from the present order of things, when there are but three families in the entire world, none of which enters North America. The Eocene family of the †titanotheres became extinct at the end of the lower substage of the White River, but in that substage there was a marvellous abundance of these huge beasts, some of which were of almost elephantine stature and bulk. The pair of great bony, horn-like protuberances on the nose varied much in size and form in the different species, short to very long, triangular, cylindrical, flattened and shovel-shaped, and gave these ungainly creatures somewhat the appearance of strange and very large rhinoceroses. The †titanotheres were a typically North American family, but sent migrants to the Old World, at least two species reaching southeastern Europe. Rhinoceroses too were extremely numerous and diversified throughout the stage and are very plainly divisible into three strongly contrasted series, which are sometimes regarded as three subdivisions of the same family and sometimes put into two separate families. One of these series, the †hyracodonts (†Hyracodon), was composed of small, long-necked and long-legged, slender and lightly built, cursorial animals, but with short, heavy heads, which gave them a somewhat clumsy look; having neither horns nor tusks, they were entirely defenceless and depended for their safety upon speed alone. The second series, or †amynodonts (†Metamynodon), formed the very antithesis of the first,—large, heavy, short-necked, and short-legged and probably amphibious in manner of life, they were armed with formidable tusks; and their skulls were so curiously modified as to bear a distinct resemblance to the skull of a huge carnivore. The †amynodonts migrated to the Old World and occur in the Oligocene of France, but the †hyracodonts would seem never to have left North America. The third series, that of the true rhinoceroses, comprised several genera at different levels in the White River beds (†Trigonias, †Cænopus, etc.); they were of uncertain origin and it has not yet been determined whether they were immigrants or of native stock. Many species have been found, varying much in size, up to that of a modern tapir, and not unlike one in proportions, for they were of lighter build and had relatively longer legs than any existing rhinoceros. The species of the lower and middle substages were all hornless, but in the uppermost substage we find skulls with a pair of nasal horns in an incipient stage of development. This was the beginning of the †paired-horned rhinoceroses (†Diceratherium) which so flourished in the John Day and the lower Miocene.

Fig. 135.—†Hornless rhinoceros (†Cænopus tridactylus) of the White River stage. Restored from a skeleton in the American Museum.

Of the horses there was no great variety and all the species so far discovered are included in a single genus (†Mesohippus), though there was a decided increment in the size of the successive species from the earlier to the later portion of the stage. Looked at superficially, it seems absurd to call these little creatures “horses” at all and the term can be justified only as implying that they were ancestral members of the family. The largest of the White River species hardly exceeded a sheep in size and all of them had comparatively short necks, long and slender legs and three-toed feet. The low-crowned grinding teeth show that they were browsers, not grazers. The abundant Eocene family of the †Lophiodontidæ made its last appearance in the White River, where it was scantily represented by slender, long-legged animals (†Colodon), with feet singularly like those of the contemporary horses, except that there were four toes in the front foot. Tapirs (†Protapirus) were very much less common than rhinoceroses or horses and were hardly half as large as the existing species of the family and of relatively far more slender form; the development of the proboscis had already begun. Lastly, the presence of the clawed †chalicotheres has been reported from the lower Oligocene of Canada, but the material is too fragmentary for generic reference.

Though the number of artiodactyl families yet identified among the White River fossils is no larger than that of the perissodactyl families, the artiodactyls greatly preponderated in individual abundance. The peccaries, which were fairly common, resembled those of the John Day, but were considerably smaller. Of the camels, there were two series, one of which (†Eotylopus), lately described by Dr. Matthew, is of yet unknown significance, while the other (†Poëbrotherium) was apparently the ancestor common to all the subsequent phyla of camels and llamas. This extremely interesting genus had species which ranged in size from a gazelle to a sheep, had two toes in each foot, a moderately elongate neck and teeth which were beginning to assume the high-crowned character. From this it may be inferred that those animals were, partly at least, of grazing habit, which was rare among White River ungulates, most of which fed upon leaves and soft and succulent plants. An extinct family, the †Hypertragulidæ, were a greatly diversified group of dainty little creatures, one of which (†Hypisodus) was no larger than a rabbit and had high-crowned teeth. The other genera (†Leptomeryx, †Hypertragulus) must have resembled in form and proportions the tiny little chevrotains or “mouse-deer” of the East Indian islands. Late in the age arose a larger form of this family, nearly equalling the Musk-Deer in size, the extraordinary genus †Protoceras, which was, especially the males, a grotesque object. The males had a pair of upper canine tusks and two pairs of prominent long protuberances on the skull. This, or some similar form, must have been the ancestor of the still more bizarre †Syndyoceras of the lower Miocene.

The †oreodonts were by far the commonest of White River mammals, and evidently they roamed the woods and plains in great herds. There were several species, larger and smaller, of the abundant genus (†Merycoidodon) but the largest did not surpass a modern peccary in size and was somewhat like that animal in appearance, but had a shorter head and much longer tail. In the upper substage appeared a very peculiar genus of this family (†Leptauchenia), animals with short, deep, almost monkey-like heads, and presumably aquatic in habits. The †agriochœrids were very much less common; they may be described roughly as †oreodonts with very long, cat-like tails and clawed feet.

Fig. 136.—†Merycoidodon culbertsoni, the most abundant of White River †oreodonts. Restored from a skeleton in the American Museum of Natural History.