The existing members of this order can be readily grouped into the Hyracoidea, Proboscidea, Perissodactyla, and Artiodactyla, each of which divisions has quite the value of an order, and all of which are sharply marked off from each other. But as the discovery of so many fossil forms has to a great extent rendered these demarcations less sharp, it is better to regard all these groups as not more than sub-orders of a larger "Order" Ungulata. Even when this conclusion has been necessarily arrived at from a consideration of the more ancient groups of Ungulate animals, the definition of such an order remains a difficult matter for the systematist. For the earliest of these forms, more particularly the Ancylopoda, the Amblypoda, and the Condylarthra, whose peculiarities will be dealt with at length subsequently, are not by any means easily differentiated from the primitive Carnivorous mammals of that date, the Creodonta; these latter, moreover, fade into the Marsupials through the so-called Sparassodonta of Professor Ameghino. To confine ourselves to the Ungulates, we may perhaps define them as terrestrial animals with hoofs rather than claws or nails, and chiefly, if not entirely, vegetarian in habit. The teeth are bunodont or lophodont, the tendency to the production of the latter type being always marked. The walk, although plantigrade in the older types, becomes more and more digitigrade, except in such survivals from antiquity as Hyrax. There is, too, as we pass from the ancient types to the modern, a gradual perfection of the limbs as running

and not climbing or grasping organs; the number of toes becomes reduced, and culminates twice (in the horse and in the Litopterna) in one toe on each foot; at the same time the ulna becomes rudimentary and fuses with the radius, and the fibula in the hind-limb undergoes a like reduction. The clavicle is absent even in some of the oldest types; its presence in Typotherium[[114]] is highly remarkable. The tail too, an organ which is long in some of the early forms, gets short in their modern derivatives.

Fig. 110.—An early Ungulate. Phenacodus primaevus. × 1⁄12. (After Osborn.)

Fig. 111.—Series of metacarpals and metatarsals of Camelidae, to show secular and progressive increase in size. From left to right the species are Protylopus petersoni, Poebrotherium labiatum, Gomphotherium sternbergi, Procamelus occidentalis. F, Fore-foot; H, hind-foot; III, IV, third and fourth metapodials. (After Wortman.)

Coupled with the increasing perfection of the foot as an organ used merely for the support of the body, certain interesting changes have taken place in the arrangement with regard to each other of the several bonelets of the wrist and ankle. It has been held by Cope and others that the truly primitive disposition of these bones was that presented to us by certain early types, such as Meniscotherium or the existing elephant or Hyrax. In these animals there is (see Fig. 112) a serial

arrangement of these bones, the distal bones only, or very nearly only, articulating with the corresponding bones in the upper series. In the modern types (cf. Fig. 113) there is, on the other hand, an interlocking, so that the bones of the distal series articulate with two of those of the proximal series. By this is produced, as it would appear, a much firmer foot, less liable to "give" under pressure, and thus more fitted for an animal that runs. It is the same principle as that adopted in the laying of bricks. The actual stress and strain of impact has been held responsible for those changes. An equally ingenious and possibly truer explanation of the undoubted facts has lately been advanced by Mr. W. D.

Matthew.[[115]] He has pointed out that in some ancient Ungulates the carpus is not serial but interlocking, even in forms which belong to the earliest Eocene groups, such as the genus Protolambda among the Amblypoda. Now in the fore-foot of Meniscotherium and the living Hyrax there is a separate centrale which is wanting in the greater number of Ungulates. The absorption, that is the practical dropping out of this bone, would restore to an interlocking carpus the serial arrangement; while on the other hand, by the fusion of this bone with the scaphoid, the interlocking disposition would be maintained.