Still commoner were the Artiodactyla. Many species of grazing camels, which were the predominant artiodactyl family in North America during upper Miocene times, were the ancestors both of the true camels of the Old World and the South American llamas. †Giraffe-camels have not yet been found and no doubt they were much less abundant than in the middle Miocene, but that they had not completely disappeared is shown by their recurrence in the Pliocene. As compared with earlier ages, the †oreodonts had begun a rapid decline and had lost notably both in numbers and variety, but one most curious beast (†Pronomotherium, [Fig. 197, p. 375]) marked the final step in the development of the short-faced, proboscis-bearing series, which may be traced back to its beginnings in the Oligocene. In this wonderful creature the skull was so short and deep as to suggest that of a gorilla or some other great ape. No other artiodactyls even approximate these later proboscis-bearing †oreodonts in the altogether exceptional form of the skull. Grazing †oreodonts (†Merychyus), of moderate and small size with high-crowned teeth, were evidently quite common on the upper Miocene plains. The †hornless deer and “†deer-antelopes” differed but little from those of the lower Pliocene. Peccaries were fairly abundant.
Fig. 126.—†Procamelus elrodi, a large camel from the upper Miocene. Restored from specimens in the Carnegie Museum.
The upper Miocene fauna was especially characterized by the large number of mammals, belonging to several different orders, which had acquired the high-crowned, persistently growing pattern of grinding teeth. Many of the horses, camels, ruminants and rodents displayed this structure, and, as was first pointed out by Kowalevsky, the explanation is probably to be found in the spread of grassy plains at the expense of the forests. On account of the silica which they contain, the grasses are very abrasive and rapidly wear the teeth down. In adaptation to this new source of abundant and nutritious food, many kinds of mammals developed a form of tooth which was fitted to compensate by growth for the loss through abrasion.
The middle Miocene, small areas of which occur in Montana, eastern Oregon and northeastern Colorado, has received various local names, the typical one being the Deep River of Montana. Very probably, these scattered areas are not strictly contemporaneous, but form a closely connected series. That a land-connection with the eastern hemisphere existed, is made clear by the appearance of several unmistakably Old World types of animals and the beginnings of migration from South America are perhaps also to be noted, though this cannot be positively stated. The evidence for the South American connection is the finding in the middle Miocene of Oregon of what are believed to be the earliest remains of †ground-sloths yet discovered in North America, but the material is too scanty for altogether certain determination.
The smaller animals are not very well represented in the middle Miocene faunas, as conditions appear to have been unfavourable to their preservation; something is known of them, nevertheless. The very curious extinct family of rodents known as the †Mylagaulidæ, the presence of which was noted in the upper Miocene and lower Pliocene, first appeared here. These †mylagaulids, which were distantly related to the modern Sewellel (Aplodontia rufa), were characterized by the great enlargement and complication of one of the grinding teeth in each jaw and the consequent reduction of the others. One genus of this family, as in the Pliocene, had the peculiarity, unique among rodents, of developing a large horn upon the nose, like a miniature rhinoceros. Among the Carnivora, we find a great variety of dogs, large and small, all belonging to extinct genera, as indeed is true of the other carnivores also. True felines have been found, but as yet, none of the †sabre-tooth series; the abundance of the latter, however, in both preceding and succeeding formations, is sufficient proof that the discovery of them in the middle Miocene is merely a question of time. Mustelines were present, and especially noteworthy is the appearance of the first American otters, immigrants from the Old World.
Of the hoofed animals, the most interesting are the Proboscidea, the most ancient of which that are definitely determinable in America occur in this horizon. The place of origin and ancestry of these animals were long exasperating puzzles. Appearing suddenly in the Miocene of Europe and North America, in which regions nothing was known that could, with any plausibility, be regarded as ancestral to them, they might as well have dropped from the moon, for all that could be told concerning their history. The exploration of the Eocene and Oligocene beds of Egypt has dispelled the mystery and shown that Africa was the original home of the group, whence they gradually spread to every continent except Australia. Little is known of these earliest American proboscideans, but they were doubtless small †mastodons of the four-tusked type.
Among the Perissodactyla, the rhinoceroses were perhaps the most conspicuous; the native American stocks of this family appear to have mostly died out and to have been replaced by two or more phyla of immigrants from the Old World, some of which were hornless, others had a small horn on the tip of the nose and others again had a second and smaller horn on the forehead. Tapirs, though unquestionably present, are rare as fossils and not well known. Several distinct phyla of horses may be distinguished, which were like small ponies in size, but of more slender form; they were all three-toed, but there were marked differences among them with regard to the degree to which the middle toe (the third of the original five) had been enlarged to carry the whole weight and the lateral toes (second and fourth) reduced to mere “dew-claws.” While browsing horses, with low-crowned teeth, still persisted in large numbers, we find also the extremely interesting beginnings of the highly complex, cement-covered and high-crowned teeth of the grazing kinds. The clawed †chalicotheres were present, though very little is known about them because of the fragmentary character of the remains.
The Artiodactyla were much more varied and abundant, though they did not rival the great assemblage of these animals found in the European Miocene. Of the peccaries little more can be said than that they were present in these faunas. The †oreodonts were very numerous, both individually and generically; two stages of the proboscis-bearing kind are found here together, the older, long-faced genus (†Promerycochœrus) surviving from the Oligocene, while the newer Miocene type was short-faced and had a moderate proboscis (see [Fig. 196, p. 373]). Others had more the proportions of peccaries and still others were very small and presumably aquatic in habits. Camels abounded, both the grazing kinds which were ancestral to the modern forms of South America and Asia, and the great, browsing †giraffe-camels. The †hornless deer and the antlered †deer-antelopes were much like those of the Upper Miocene, slender and graceful little creatures, and there were also considerably larger ruminants (†Dromomeryx) with straight, simple and non-deciduous horns, which may be called antelopes.