The very extraordinary and aberrant animals which are referable to this suborder have been understood only since the year 1888, for, as was shown in an earlier chapter ([p. 41]) their scattered parts had been assigned to two different mammalian orders, the skull to the perissodactyls and the feet to the pangolins, or scaly anteaters (Pholidota) of the Old World, since it occurred to no one that the same animal could have such a skull and teeth in combination with such feet.
The history of the Ancylopoda is still very incomplete, only four genera, of the lower Pliocene, middle and lower Miocene, and the middle Eocene respectively, being at all adequately known, but even in this imperfect form the story is worth telling. The suborder was probably of American origin and its most ancient known member existed in the middle Eocene. Both in Europe and North America the group persisted into the lower Pliocene and it is believed, though not clearly demonstrated, that in eastern Asia it continued even into the Pleistocene. All the genera of the suborder may be included in a single family.
7. †Chalicotheriidæ. †Chalicotheres
The specimens which so far have been found in the American middle and upper Miocene and lower Pliocene are very fragmentary, consisting of little more than teeth, and give no information other than to demonstrate the presence of the family in North America during that period of time. On the other hand, the European genera of the middle Miocene and lower Pliocene are well known and may or may not have been closely similar to their American contemporaries, though they were undoubtedly larger. In these most peculiar and grotesque animals (†Macrotherium and †Chalicotherium) the head was relatively small, the teeth were very low-crowned and adapted only to a diet of soft vegetable substances and the mode of feeding must have been that of browsing upon leaves and shoots of trees and bushes; the premolars had not acquired the molar-pattern, which was very exceptional for perissodactyls of so late a time, such a difference between the two classes of teeth being characteristic of the Eocene members of the order; the incisors and canines were reduced, but the formula is not definitely known.
The neck was of moderate length, the body very long, and the limbs were also elongate, especially the anterior pair, in consequence of which the back sloped downward from the shoulders to the rump; the two fore-arm bones were fused together, and these, with the thigh-bones, were the longest segments of the limbs. The special peculiarity of these animals was in the character of the feet, which had three toes, each armed with a huge claw, instead of terminating in a hoof, as it does in all normal perissodactyls. The external digit, which, in the absence of the fifth, was the fourth, was the largest of the series and apparently bore the most of the weight, a notable departure from the normal perissodactyl symmetry, in which the third or median toe is the largest. The hind feet were considerably smaller than the fore, but had similar claws.
Many suggestions have been offered as to the manner in which these great claws were employed. The teeth demonstrate that these animals could not have had predaceous habits, but must have been inoffensive plant-feeders. As no such herbivorous creatures are living now, it is impossible to reach a definitive solution of the problem, which is further complicated by the fact that in two other orders of hoofed mammals, Artiodactyla and †Toxodontia, a more or less similar transformation of hoofs into claws took place, and among the edentates the large, herbivorous †ground-sloths (†Gravigrada) had enormous claws. It is inadmissible to suppose that these great †chalicotheres could have been burrowers, or tree-climbers, or that they pursued and slaughtered prey of any kind, for, aside from the character of the teeth, such heavy and slow-moving beasts would have been utterly inefficient at work of that sort. No doubt, the claws were used, to some extent, as weapons of defence, as the existing South American Ant-Bear (Myrmecophaga jubata) uses his formidable claws; probably also some, if not all, of these clawed ungulates would employ the fore feet in digging for roots and tubers, as is done by the bears generally. Many years ago, the late Sir Richard Owen suggested with reference to the †ground-sloths that the principal use of the fore feet, other than that of locomotion, was to draw down within reach of the long tongue and prehensile upper lip the branches upon which they browsed. This explanation may perhaps be applicable to all of these aberrant and exceptional groups of hoofed animals.
Fig. 185.—Left manus of lower Miocene †chalicothere (†Moropus). (After Peterson.)
In the lower Miocene (Arikaree stage) of North America well-nigh complete skeletons of a large †chalicothere (†Moropus, [Fig. 130, p. 240]) have been obtained, an animal which considerably exceeded a large horse in bulk and stature. In structure this genus had departed less widely from the normal perissodactyl type than the genera of the European Miocene and Pliocene above described and was in many respects more primitive. It could not, however, have been directly ancestral to the European forms, though indicating in a general sort of way what the ancestral type must have been. †Moropus had a relatively small, slender and pointed head, a long neck, much longer than in the European genera, and long fore legs; the shorter hind legs gave the back a steep inclination from the shoulders to the rump. The proportions of the head, neck and limbs suggest those of a giraffe, in less exaggerated form, but the likeness is more marked in the skeleton than in the restoration and is at best a distant one. The feet were armed with the great claws characteristic of the suborder, but the fore foot, in addition to the three functional toes, had a long splint, representing the rudimentary fifth digit; of the first, or pollex, no trace remained. The perissodactyl plan of symmetry had not yet been lost, the third or median digit being the longest of the series. In the hind foot, which had only three toes, the departure from the perissodactyl arrangement had already begun, and the third and fourth digits (i.e. of the original five) were of nearly equal size, both in length and thickness, while the second was smaller.
The family is represented in the John Day, or upper Oligocene, by specimens which are sufficiently characteristic to prove that they are properly referable to this group. They have been assigned to the same genus as that of the lower Miocene, but whether the identification is justified remains to be determined.