Part 8

I have said that the structure of the grinding teeth is very complicated, the harder and the softer parts being, as it were, interlaced with one another. The result of this is that, as the tooth wears, the crown presents a peculiar pattern, the nature of which is not very easily deciphered at first, but which it is important we should understand clearly. Each grinding tooth of the upper jaw has an outer wall so shaped that, on the worn crown, it exhibits the form of two crescents, one in front and one behind, with their concave sides turned outwards. From the inner side of the front crescent, a crescentic front ridge passes inwards and backwards, and its inner face enlarges into a strong longitudinal fold or pillar. From the front part of the hinder crescent, a back ridge takes a like direction, and also has its pillar.

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.

The ulna is slightly more distinct than in the horse; and the whole length of it, as a very slender shaft intimately united with the radius, is completely traceable. The fibula appears to be in the same condition as in the horse. The teeth of the Hipparion are essentially similar to those of the horse, but the pattern of the grinders is in some respects a little more complex, and there is a depression on the face of the skull in front of the orbit, which is not seen in existing horses.

In the earlier Miocene, and perhaps the later Eocene deposits of some parts of Europe, another extinct animal has been discovered, which Cuvier, who first described some fragments of it, considered to be a Palæotherium. But as further discoveries threw new light on its structure, it was recognized as a distinct genus under the name of Anchitherium.

In its general characters, the skeleton of Anchitherium is very similar to that of the horse. In fact, Lartet and De Blainville called it Palæotherium equinum or hippoides; and De Christol, in 1847, said that it differed from Hipparion in little more than the characters of its teeth, and gave it the name of Hipparitherium. Each foot possesses three complete toes; while the lateral toes are much larger in proportion to the middle toe than in Hipparion, and doubtless rested on the ground in ordinary locomotion.

The ulna is complete and quite distinct from that radius, though firmly united with the latter. The fibula seems also to have been complete. Its lower end, though intimately united with that of the tibia, is clearly marked off from the latter bone.

There are forty-four teeth. The incisors have no strong pit. The canines seem to have been well developed in both sexes. The first of the seven grinders, which, as I have said, is frequently absent, and when it does exist, is small in the horse, is a good-sized and permanent tooth, while the grinder which follows it is but little larger than the hinder ones. The crowns of the grinders are short, and though the fundamental pattern of the horse-tooth is discernible, the front and back ridges are less curved, the accessory pillars, are wanting, and the valleys, much shallower, are not filled up with cement.

Seven years ago, when I happened to be looking critically into the bearing of palæontological facts upon the doctrine of evolution, it appeared to me that the Anchitherium, the Hipparion, and the modern horses, constitute a series in which the modifications of structure coincide with the order of chronological occurrence, in the manner in which they must coincide, if the modern horses really are the result of the gradual metamorphosis, in the course of the Tertiary epoch, of a less specialized ancestral form. And I found by correspondence with the late eminent French anatomist and palæontologist, M. Lartet, that he had arrived at the same conclusion from the same data.

That the Anchitherium type had become metamorphosed into the Hipparion type, and the latter into the Equine type, in the course of that period of time which is represented by the latter half of the Tertiary deposits, seemed to me to be the only explanation of the facts for which there was even a shadow of probability.

And, hence, I have ever since held that these facts afford evidence of the occurrence of evolution, which, in the sense already defined, may be termed demonstrative.

All who have occupied themselves with the structure of Anchitherium, from Cuvier onwards, have acknowledged its many points of likeness to a well-known genus of extinct Eocene mammals, Palæotherium. Indeed, as we have seen, Cuvier regarded his remains of Anchitherium as those of a species of Palæotherium. Hence, in attempting to trace the pedigree of the horse beyond the Miocene epoch and the Anchitheroid form, I naturally sought among the various species of Palæotheroid animals for its nearest ally, and I was led to the conclusion that the Palæotherium minus (Plagiolophus) represented the next step more nearly than any form then known.

I think that this opinion was fully justifiable; but the progress of investigation has thrown an unexpected light on the question, and has brought us much nearer than could have been anticipated to a knowledge of the true series of the progenitors of the horse.

You are all aware that, when your country was first discovered by Europeans, there were no traces of the existence of the horse on any part of the American Continent. The accounts of the conquest of Mexico dwell upon the astonishment of the natives of that country when they first became acquainted with that astounding phenomenon--a man seated upon a horse. Nevertheless, the investigations of American geologists have proved that the remains of horses occur in the most superficial deposits of both North and South America, just as they do in Europe. Therefore, for some reason or other--no feasible suggestion on that subject, so far as I know, has been made--the horse must have died out on this continent at some period preceding the discovery of America. Of late years there has been discovered in your Western Territories that marvellous accumulation of deposits, admirably adapted for the preservation of organic remains, to which I referred the other evening, and which furnishes us with a consecutive series of records of the fauna of the older half of the Tertiary epoch, for which we have no parallel in Europe. They have yielded fossils in an excellent state of conservation and in unexampled numbers and variety. The researches of Leidy and others have shown that forms allied to the Hipparion and the Anchitherium are to be found among these remains. But it is only recently that the admirably conceived and most thoroughly and patiently worked-out investigations of Professor Marsh have given us a just idea of the vast fossil wealth, and of the scientific importance, of these deposits. I have had the advantage of glancing over the collections in Yale Museum; and I can truly say, that so far as my knowledge extends, there is no collection from any one region and series of strata comparable, for extent, or for the care with which the remains have been got together, or for their scientific importance, to the series of fossils which he has deposited there. This vast collection has yielded evidence bearing upon the question of the pedigree of the horse of the most striking character. It tends to show that we must look to America, rather than to Europe, for the original seat of the equine series; and that the archaic forms and successive modifications of the horse's ancestry are far better preserved here than in Europe.

Professor Marsh's kindness has enabled me to put before you a diagram, every figure of which is an actual representation of some specimen which is to be seen at Yale at this present time (Fig. 9).

The succession of forms which he has brought together carries us from the top to the bottom of the Tertiaries. Firstly, there is the true horse. Next we have the American Pliocene form of the horse (Pliohippus); in the conformation of its limbs it presents some very slight deviations from the ordinary horse, and the crowns of the grinding teeth are shorter. Then comes the Protohippus, which represents the European Hipparion, having one large digit and two small ones on each foot, and the general characters of the fore-arm and leg to which I have referred. But it is more valuable than the European Hipparion for the reason that it is devoid of some of the peculiarities of that form--peculiarities which tend to show that the European Hipparion is rather a member of a collateral branch, than a form in the direct line of succession. Next, in the backward order in time, is the Miohippus, which corresponds pretty nearly with the Anchitherium of Europe. It presents three complete toes--one large median and two smaller lateral ones; and there is a rudiment of that digit, which answers to the little finger of the human hand.

The European record of the pedigree of the horse stops here; in the American Tertiaries, on the contrary, the series of ancestral equine forms is continued into the Eocene formations. An older Miocene form, termed Mesohippus, has three toes in front, with a large splint-like rudiment representing the little finger; and three toes behind. The radius and ulna, the tibia and the fibula, are distinct, and the short crowned molar teeth are anchitheroid in pattern.

But the most important discovery of all is the Orohippus, which comes from the Eocene formation, and which is the oldest member of the equine series, as yet known. Here we find four complete toes on the front-limb, three toes on the hind-limb, a well-developed ulna, a well-developed fibula, and short-crowned grinders of simple pattern.

Thus, thanks to these important researches, it has become evident that, so far as our present knowledge extends, the history of the horse-type is exactly and precisely that which could have been predicted from a knowledge of the principles of evolution. And the knowledge we now possess justifies us completely in the anticipation, that when the still lower Eocene deposits, and those which belong to the Cretaceous epoch, have yielded up their remains of ancestral equine animals, we shall find, first, a form with four complete toes and a rudiment of the innermost or first digit in front, with probably, a rudiment of the fifth digit in the hind foot; while, in still older forms, the series of the digits will be more and more complete, until we come to the five-toed animals, in which, if the doctrine of evolution is well founded, the whole series must have taken its origin.

That is what I mean by demonstrative evidence of evolution. An inductive hypothesis is said to be demonstrated when the facts are shown to be in entire accordance with it. If that is not scientific proof, there are no merely inductive conclusions which can be said to be proved. And the doctrine of evolution, at the present time, rests upon exactly as secure a foundation as the Copernican theory of the motions of the heavenly bodies did at the time of its promulgation. Its logical basis is precisely of the same character--the coincidence of the observed facts with theoretical requirements.