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CHAPTER V OF LITTLE MASTERPIECES OF SCIENCE THE NATURALIST AS INTERPRETER AND SEER All LibriVox recordings are in the public domain. For more information or to volunteer, please visit LibriVox.org. Read by Paul Lawley-Jones

THE NATURALIST AS INTERPRETER AND SEER Edited by George Isles THE EVOLUTION OF THE HORSE Thomas Henry Huxley Professor Huxley, as a naturalist, educator and controversialist, was one of the commanding figures of the 19th century.

To physiology and morphology, his researches added much of importance. As an expositor, he stood unapproached. As the bold and witty champion of Darwinism, he gave natural selection and acceptance much more early and wide than it would otherwise have enjoyed. In 1876, he delivered in America three lectures on evolution. The third of the series is here given.

All three are copyrighted and published by D. Appleton & Co. New York, in a volume which also contains a lecture on the study of biology. Since 1876, the arguments of Professor Huxley have been reinforced by the discovery of many fossils connecting not only the horse, but other quadrupeds, with species widely different and now extinct. The most comprehensive collection illustrating the descent of the horse

is to be seen at the American Museum of Natural History, New York, where also the evolution of tapirs, camels, llamas, rhinoceroses, dinosaurs, great ground sloths and other animals are clearly to be traced, in most cases by remains discovered in America. A capital book on the theme brooch by Professor Huxley is Animals of the Past

by Frederick A. Lucas, curator of the Division of Comparative Anatomy, United States National Museum, Washington, D.C., published by McClure, Phillips & Co., New York. The Life and Letters of Professor Huxley, edited by his son Leonard Huxley, is a work of rare interest. It is published by D. Appleton & Co., New York. The occurrence of historical facts is said to be demonstrated when the evidence that they happened

is of such a character as to render the assumption that they did not happen in the highest degree improbable. And the question I have now to deal with is whether evidence in favour of the evolution of animals of this degree of cogency is, or is not, obtainable from the record of the succession of living forms which is presented to us by fossil remains. Those who have attended to the progress of paleontology are aware that evidence of the character which I have defined has been produced in considerable and continually increasing quantity during the last few years. Indeed, the amount and the satisfactory nature of that evidence are somewhat surprising when we consider the conditions under which alone we can hope to obtain it

It is obviously useless to seek for such evidence except in localities in which the physical conditions have been such as to permit of the deposit of an unbroken, or but rarely interrupted series of strata through a long period of time in which the group of animals to be investigated has existed in such abundance

as to furnish the requisite supply of remains, and in which, finally, the materials composing the strata are such as to ensure the preservation of these remains in a tolerably perfect and undisturbed state. It so happens that the case which, at present,

most nearly fulfills all these conditions is that of the series of extinct animals which culminates in the horses, by which term I mean to denote not merely the domestic animals with which we are all so well acquainted, but their allies, the ass, zebra, quagga and the like.

In short, I use horses as the equivalent of the technical name equidae which is applied to the whole group of existing equine animals. The horse is in many ways a remarkable animal, not least so in the fact that it presents us with an example of one of the most perfect pieces of machinery in the living world.

In truth, among the works of human ingenuity it cannot be said that there is any locomotive so perfectly adapted to its purposes, doing so much work with so small a quantity of fuel as this machine of nature's manufacture, the horse. And, as a necessary consequence of any sort of perfection, of mechanical perfection as of others, you find that the horse is a beautiful creature, one of the most beautiful of all land

animals. Look at the perfect balance of its form and the rhythm and force of its action. The locomotive machinery is, as you are aware, resident in its slender fore and hind limbs. They are flexible and elastic levers, capable of being moved by very powerful muscles, and in order to supply the engines which work these levers with the force which they expend, the horse is provided with a very perfect apparatus for grinding its food and extracting therefrom the requisite fuel.

Without attempting to take you very far into the region of osteological detail, I must nevertheless trouble you with some statements respecting the anatomical structure of the horse. And more especially, will it be needful to obtain a general conception of the structure of its fore and hind limbs and of its teeth. But I shall only touch upon these points which are absolutely essential to our inquiry. Let us turn in the first place to the forelimb.

In most quadrupeds, as in ourselves, the forearms contain distinct bones called the radius and the ulna. The corresponding region in the horse seems at first to possess but one bone Careful observation, however, enables us to distinguish in this bone a part which clearly answers to the upper end of the ulna This is closely united with the chief mass of the bone which represents the radius

and runs out into a slender shaft which may be traced for some distance downwards upon the back of the radius and then in most cases thins out and vanishes It takes still more trouble to make sure of what is nevertheless the fact that a small part of the lower end of the bone of the horse's forearm

which is only distinct in a very young foal is really the lower extremity of the ulna What is commonly called the knee of a horse is its wrist The cannon bone answers to the middle bone of the five metacarpal bones which support the palm of the hand in ourselves

The pastern, coronary and coffin bones of the veterinarians answer to the joints of our middle fingers, while the hoof is simply a greatly enlarged and thickened nail. But if what lies below the horse's knee thus corresponds to the middle finger in ourselves, what has become of the four other fingers or digits? We find in the places of the second and fourth digits only two slender splint-like bones,

about two-thirds as long as the cannon bone, which gradually taper to their lower ends and bear no finger joints, or, as they are termed, phalanges. Sometimes small bony or grisly nodules are to be found at the bases of these two metacarpal splints, and it is probable that these represent rudiments of the first and fifth toes. Thus, the part of the horse's skeleton which corresponds with that of the human hand, contains one overgrown middle digit and at least two

imperfect lateral digits, and these answer respectively to the third, the second, and the fourth fingers in man. Corresponding modifications are found in the hind limb. In ourselves, and in most quadrupeds, the leg contains two distinct bones, a large bone, the tibia, and a smaller and more slender bone, the fibula. But in the horse, the fibula seems, at first, to be reduced to his upper end, a short slender bone united with the tibia and ending in a point below, occupying its

place. Examination of the lower end of a young foal's shin bone, however, shows a distinct portion of osseous matter, which is the lower end of the fibula, so that the apparently single lower end of the shin bone is really made up of the coalesced ends of the tibia and fibula just as the apparently single lower end of the forearm bone is composed of the coalesced radius and ulna. The heel of the horse is the part commonly known as the hock. The hinder cannon bone enters to the

middle metatarsal bone of the human foot, the pastern, coronary and coffin bones to the middle toe bones, the hind hoof to the nail, as in the forefoot. And, as in the forefoot, there are merely two splints to represent the second and fourth toes. Sometimes a rudiment of a fifth toe appears to be traceable. The teeth of a horse are not less peculiar than its limbs. The living engine, like all others, must be well stoked if it is to do its work, and the horse, if it is to make good

its wear and tear and to exert the enormous amount of force required for its propulsion must be well and rapidly fed. To this end, good cutting instruments and powerful and lasting crushers are needful. Accordingly, the 12 cutting teeth of a horse are close set and concentrated in the forepart of its mouth, like so many adzes or chisels. The grinders or molars are large and have an extremely complicated structure, being composed of a number of different substances of

unequal hardness. The consequence of this is that they wear away at different rates, and, hence, the surface of each grinder is always as uneven as that of a good millstone. 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, embedded 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 anyone 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 44 While in horses, the usual number is 40, and in the absence of the canines, it may be reduced to 36 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 forearm and of the leg complete and separate,

and which possessed 44 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 forearm 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 Quartinari 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 the 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 dewclaws such as are to be found in many ruminant animals. The hipparian, as the extinct European three-toed horse is called, in fact presents a foot similar to that of the American proto-hippus, except that in the hipparian the smaller digits are situated farther back

and are of smaller proportional size than in the proto-hippus. 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 Hipparian 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 paleotherium. 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, Latte and de Blainville called it Paleotherium equinum, or Hippoides. and de Cristal in 1847 said that it differed from hipparian in little more than the characters of its teeth and gave it the name of hipparotherium. Each foot possesses three complete toes

while the lateral toes are much larger in proportion to the middle toe than in hipparian 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 44 teeth The incisors have no strong pit The canine 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 paleontological facts upon the doctrine of evolution, it appeared to me that the Ancetherium, the Hipparian 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 paleontologist Monsieur Latte that he had arrived at the same conclusion from the same data. That the Ancetherium type had become metamorphosed into the Hipparian 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 paleotherium indeed as we have seen

cuvier regarded his remains of anchitherium as those of a species of paleotherium hence in attempting to trace the pedigree of the horse beyond the Miocene epoch and the architheroid form, I naturally sought among the various species of paleotheroid animals for its nearest ally, and I was led to the conclusion that the Paleotherium 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 account 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 marvelous 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 researchers of Ledy and others have shown that forms allied to the Hipparian and the Architherium 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 collection 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 has been 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. 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 Pliocenean form of the horse, Plia hippus.

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 proto-hippus, which represents the European Hipparion, having one large digit and two small ones on each foot, and the general characters of the forearm 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 Myohippus, which corresponds pretty nearly with the Anchetherium 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 fibula, are distinct

and the short-crowned molar teeth are anchethyroid 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 crown 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 toad 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 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 the only way of escape if it be a way of escape from the conclusions which i have just indicated is the supposition that all these different equine forms have been created separately at separate epochs of time. And, I repeat, that of such an hypothesis as this

there neither is, nor can be, any scientific evidence. And assuredly, so far as I know, there is none which is supported, or pretends to be supported, by evidence or authority of any other kind. I can but think that the time will come when suggestions such as these, such obvious attempts to escape the force of demonstration, will be put upon the same footing as the supposition made by some writers who are, I believe, not completely extinct at present,

that fossils are mere simulacra, images, are no indications of the former existence of the animals to which they seem to belong, but that they are either sports of nature or special creations intended, as I heard suggested the other day, to test our faith. In fact, the whole evidence is in favour of evolution, and there is none against it. And I say this although perfectly well aware of the seeming difficulties which have been built up upon what appears

to the uninformed to be a solid foundation. I meet constantly with the argument that the doctrine of evolution cannot be well founded because it requires the lapse of a very vast period of time, while the duration of life upon the earth, thus implied, is inconsistent with the conclusions arrived at by the astronomer and the physicist. I may venture to say that I am familiar with those conclusions. Inasmuch as some years ago, when President of the Geological

Society of London, I took the liberty of criticising them and of showing in what respects, as it appeared to me, they lacked complete and thorough demonstration. But putting that point aside, suppose that, as the astronomers, or some of them, and some physical philosophers tell us, It is impossible that life could have endured upon the earth for so long a period as is required by the doctrine of evolution, supposing that to be proved. I desire to be informed, what is the foundation for the statement that evolution does require so great a time?

The biologist knows nothing whatever of the amount of time which may be required for the process of evolution. It is a matter of fact that the equine forms, which I have described to you, occur in the order stated in the tertiary formations. But I have not the slightest means of guessing whether it took a million of years, or ten millions, or a hundred millions, or a thousand millions of years, to give rise to that series of changes. A biologist has no means of arriving at any conclusions as to the amount of time which may be needed for a certain quantity of organic change. he takes his time from the geologist the geologist considering the rate at which deposits are formed

and the rate at which denudation goes on upon the surface of the earth arrives at more or less justifiable conclusions as to the time which is required for the deposits of a certain thickness of rocks and if he tells me that the tertiary formation required 500 million years for their deposit, I suppose he has good ground for what he says, and I take that as a measure of the duration of the evolution of the horse from the Orahippus up to its present condition. And, if he is right,

undoubtedly evolution is a very slow process and requires a great deal of time. But suppose now that an astronomer or physicist, for instance my friend Sir William Thompson, tells me that my geological authority is quite wrong, and that he has weighty evidence to show that life could not possibly have existed upon the surface of the earth 500 million years ago, because the earth would have then been too hot to allow of life, my reply is, that is not my affair, settle that with

the geologist, and when you have come to an agreement among yourselves I will adopt your conclusions. We take our time from the geologists and physicists, and it is monstrous that, having taken our time from the physical philosopher's clock, the physical philosopher should turn around upon us and say we are too fast or too slow. What we desire to know is, is it a fact that evolution took place? As to the amount of time which evolution may have occupied,

we are in the hands of the physicist and the astronomer, whose business it is to deal with those questions. End of chapter 5