The deep interspaces or valleys between these ridges and the outer wall are filled by bony substance, which is called cement, and coats the whole tooth.
The pattern of the worn face of each grinding tooth of the lower jaw is quite different. It appears to be formed of two crescent-shaped ridges, the convexities of which are turned outwards. The free extremity of each crescent has a pillar, and there is a large double pillar where the two crescents meet. The whole structure is, as it were, imbedded in cement, which fills up the valleys, as in the upper grinders.
If the grinding faces of an upper and of a lower molar of the same side are applied together, it will be seen that the opposed ridges are nowhere parallel, but that they frequently cross; and that thus, in the act of mastication, a hard surface in the one is constantly applied to a soft surface in the other, and vice versa. They thus constitute a grinding apparatus of great efficiency, and one which is repaired as fast as it wears, owing to the long-continued growth of the teeth.
Some other peculiarities of the dentition of the horse must be noticed, as they bear upon what I shall have to say by and by. Thus the crowns of the cutting teeth have a peculiar deep pit, which gives rise to the well-known “mark” of the horse. There is a large space between the outer incisors and the front grinders. In this space the adult male horse presents, near the incisors on each side, above and below, a canine or “tush,” which is commonly absent in mares. In a young horse, moreover, there is not unfrequently to be seen, in front of the first grinder, a very small tooth, which soon falls out. If this small tooth be counted as one, it will be found that there are seven teeth behind the canine on each side; namely, the small tooth in question, and the six great grinders, among which, by an unusual peculiarity, the foremost tooth is rather larger than those which follow it.
I have now enumerated those characteristic structures of the horse which are of most importance for the purpose we have in view.
To any one who is acquainted with the morphology [comparative forms] of vertebrated animals, they show that the horse deviates widely from the general structure of mammals; and that the horse type is, in many respects, an extreme modification of the general mammalian plan. The least modified mammals, in fact, have the radius and ulna, the tibia and fibula, distinct and separate. They have five distinct and complete digits on each foot, and no one of these digits is very much larger than the rest. Moreover, in the least modified mammals the total number of the teeth is very generally forty-four, while in horses the usual number is forty, and in the absence of the canines it may be reduced to thirty-six; the incisor teeth are devoid of the fold seen in those of the horse: the grinders regularly diminish in size from the middle of the series to its front end; while their crowns are short, early attain their full length, and exhibit simple ridges or tubercles, in place of the complex foldings of the horse's grinders.
Hence the general principles of the hypothesis of evolution lead to the conclusion that the horse must have been derived from some quadruped which possessed five complete digits on each foot; which had the bones of the fore-arm and of the leg complete and separate; and which possessed forty-four teeth, among which the crowns of the incisors and grinders had a simple structure; while the latter gradually increased in size from before backwards, at any rate in the anterior part of the series, and had short crowns.
And if the horse has been thus evolved, and the remains of the different stages of its evolution have been preserved, they ought to present us with a series of forms in which the number of the digits becomes reduced; the bones of the fore-arm and leg gradually take on the equine condition; and the form and arrangement of the teeth successively approximate to those which obtain in existing horses.
Let us turn to the facts, and see how far they fulfil these requirements of the doctrine of evolution.
In Europe abundant remains of horses are found in the Quaternary and later Tertiary strata as far as the Pliocene formation. But these horses, which are so common in the cave-deposits and in the gravels of Europe, are in all essential respects like existing horses. And that is true of all the horses of the latter part of the Pliocene epoch. But in deposits which belong to the earlier Pliocene and later Miocene epochs, and which occur in Britain, in France, in Germany, in Greece, in India, we find animals which are extremely like horses—which, in fact, are so similar to horses that you may follow descriptions given in works upon the anatomy of the horse upon the skeletons of these animals—but which differ in some important particulars. For example, the structure of their fore and hind limbs is somewhat different. The bones which, in the horse, are represented by two splints, imperfect below, are as long as the middle metacarpal and metatarsal bones; and attached to the extremity of each is a digit with three joints of the same general character as those of the middle digit, only very much smaller. These small digits are so disposed that they could have had but very little functional importance, and they must have been rather of the nature of the dew-claws, such as are to be found in many ruminant animals. The Hipparion, as the extinct European three-toed horse is called, in fact, presents a foot similar to that of the American Protohippus ([Fig. 9]), except that in the Hipparion the smaller digits are situated farther back and are of smaller proportional size than in the Protohippus.