Chapter 4

Apart from the few fragmentary remains from the English greensand, to which I have referred, the Mesozoic rocks, older than those in which _Hesperornis_ and _Ichthyornis_ have been discovered, have afforded no certain evidence of birds, with the remarkable exception of the Solenhofen slates. These so-called slates are composed of a fine grained calcareous mud which has hardened into lithographic stone, and in which organic remains are almost as well preserved as they would be if they had been imbedded in so much plaster of Paris. They have yielded the _Archaeopteryx,_ the existence of which was first made known by the finding of a fossil feather, or rather of the impression of one. It is wonderful enough that such a perishable thing as a feather, and nothing more, should be discovered; yet, for a long time, nothing was known of this bird except its feather. But by and by a solitary skeleton was discovered which is now in the British Museum. The skull of this solitary specimen is unfortunately wanting, and it is therefore uncertain whether the _Archaeopteryx_ possessed teeth or not. [2] But the remainder of the skeleton is so well preserved as to leave no doubt respecting the main features of the animal, which are very singular. The feet are not only altogether bird-like, but have the special characters of the feet of perching birds, while the body had a clothing of true feathers. Nevertheless, in some other respects, _Archaeopteryx_ is unlike a bird and like a reptile. There is a long tail composed of many vertebrae. The structure of the wing differs in some very remarkable respects from that which it presents in a true bird. In the latter, the end of the wing answers to the thumb and two fingers of my hand; but the metacarpal bones, or those which answer to the bones of the fingers which lie in the palm of the hand, are fused together into one mass; and the whole apparatus, except the last joints of the thumb, is bound up in a sheath of integument, while the edge of the hand carries the principal quill-feathers. In the _Archaeopteryx,_ the upper-arm bone is like that of a bird; and the two bones of the forearm are more or less like those of a bird, but the fingers are not bound together--they are free. What their number may have been is uncertain; but several, if not all, of them were terminated by strong curved claws, not like such as are sometimes found in birds, but such as reptiles possess; so that, in the _Archaeopteryx,_ we have an animal which, to a certain extent, occupies a midway place between a bird and a reptile. It is a bird so far as its foot and sundry other parts of its skeleton are concerned; it is essentially and thoroughly a bird by its feathers; but it is much more properly a reptile in the fact that the region which represents the hand has separate bones, with claws resembling those which terminate the forelimb of a reptile. Moreover, it has a long reptile-like tail with a fringe of feathers on each side; while, in all true birds hitherto known, the tail is relatively short, and the vertebrae which constitute its skeleton are generally peculiarly modified.

Like the _Anoplotherium_ and the _Palaeotherium,_ therefore, _Archaeopteryx_ tends to fill up the interval between groups which, in the existing world, are widely separated, and to destroy the value of the definitions of zoological groups based upon our knowledge of existing forms. And such cases as these constitute evidence in favour of evolution, in so far as they prove that, in former periods of the world's history, there were animals which overstepped the bounds of existing groups, and tended to merge them into larger assemblages. They show that animal organisation is more flexible than our knowledge of recent forms might have led us to believe; and that many structural permutations and combinations, of which the present world gives us no indication, may nevertheless have existed.

But it by no means follows, because the _Palaeotherium_ has much in common with the horse, on the one hand, and with the rhinoceros on the other, that it is the intermediate form through which rhinoceroses have passed to become horses, or _vice versa;_ on the contrary, any such supposition would certainly be erroneous. Nor do I think it likely that the transition from the reptile to the bird has been effected by such a form as _Archaeopteryx._ And it is convenient to distinguish these intermediate forms between two groups, which do not represent the actual passage from the one group to the other, as _intercalary_ types, from those _linear_ types which, more or less approximately, indicate the nature of the steps by which the transition from one group to the other was effected.

I conceive that such linear forms, constituting a series of natural gradations between the reptile and the bird, and enabling us to understand the manner in which the reptilian has been metamorphosed into the bird type, are really to be found among a group of ancient and extinct terrestrial reptiles known as the _Ornithoscelida._ The remains of these animals occur throughout the series of mesozoic formations, from the Trias to the Chalk, and there are indications of their existence even in the later Palaeozoic strata.

Most of these reptiles, at present known, are of great size, some having attained a length of forty feet or perhaps more. The majority resembled lizards and crocodiles in their general form, and many of them were, like crocodiles, protected by an armour of heavy bony plates. But, in others, the hind limbs elongate and the fore limbs shorten, until their relative proportions approach those which are observed in the short-winged, flightless, ostrich tribe among birds.

The skull is relatively light, and in some cases the jaws, though bearing teeth, are beak-like at their extremities and appear to have been enveloped in a horny sheath. In the part of the vertebral column which lies between the haunch bones and is called the sacrum, a number of vertebrae may unite together into one whole, and in this respect, as in some details of its structure, the sacrum of these reptiles approaches that of birds.

But it is in the structure of the pelvis and of the hind limb that some of these ancient reptiles present the most remarkable approximation to birds, and clearly indicate the way by which the most specialised and characteristic features of the bird may have been evolved from the corresponding parts of the reptile.

In Fig. 6, the pelvis and hind limbs of a crocodile, a three-toed bird, and an ornithoscelidan are represented side by side; and, for facility of comparison, in corresponding positions; but it must be recollected that, while the position of the bird's limb is natural, that of the crocodile is not so. In the bird, the thigh bone lies close to the body, and the metatarsal bones of the foot (ii., iii., iv., Fig. 6) are, ordinarily, raised into a more or less vertical position; in the crocodile, the thigh bone stands out at an angle from the body, and the metatarsal bones (i., ii., iii., iv., Fig. 6) lie flat on the ground. Hence, in the crocodile, the body usually lies squat between the legs, while, in the bird, it is raised upon the hind legs, as upon pillars.

In the crocodile, the pelvis is obviously composed of three bones on each side: the ilium (_Il._), the pubis (_Pb._), and the ischium (_Is._). In the adult bird there appears to be but one bone on each side. The examination of the pelvis of a chick, however, shows that each half is made up of three bones, which answer to those which remain distinct throughout life in the crocodile. There is, therefore, a fundamental identity of plan in the construction of the pelvis of both bird and reptile; though the difference in form, relative size, and direction of the corresponding bones in the two cases are very great.

But the most striking contrast between the two lies in the bones of the leg and of that part of the foot termed the tarsus, which follows upon the leg. In the crocodile, the fibula (_F_) is relatively large and its lower end is complete. The tibia (_T_) has no marked crest at its upper end, and its lower end is narrow and not pulley-shaped. There are two rows of separate tarsal bones (_As., Ca., &c._) and four distinct metatarsal bones, with a rudiment of a fifth.

In the bird, the fibula is small and its lower end diminishes to a point. The tibia has a strong crest at its upper end and its lower extremity passes into a broad pulley. There seem at first to be no tarsal bones; and only one bone, divided at the end into three heads for the three toes which are attached to it, appears in the place of the metatarsus.

In the young bird, however, the pulley-shaped apparent end of the tibia is a distinct bone, which represents the bones marked _As., Ca.,_ in the crocodile; while the apparently single metatarsal bone consists of three bones, which early unite with one another and with an additional bone, which represents the lower row of bones in the tarsus of the crocodile.

In other words, it can be shown by the study of development that the bird's pelvis and hind limb are simply extreme modifications of the same fundamental plan as that upon which these parts are modelled in reptiles.

On comparing the pelvis and hind limb of the ornithoscelidan with that of the crocodile, on the one side, and that of the bird, on the other (Fig. 6), it is obvious that it represents a middle term between the two. The pelvic bones approach the form of those of the birds, and the direction of the pubis and ischium is nearly that which is characteristic of birds; the thigh bone, from the direction of its head, must have lain close to the body; the tibia has a great crest; and, immovably fitted on to its lower end, there is a pulley-shaped bone, like that of the bird, but remaining distinct. The lower end of the fibula is much more slender, proportionally, than in the crocodile. The metatarsal bones have such a form that they fit together immovably, though they do not enter into bony union; the third toe is, as in the bird, longest and strongest. In fact, the ornithoscelidan limb is comparable to that of an unhatched chick.

Fig. 6.--Bird. Ornithoscelidan. Crocodile. The letters have the same signification in all the figures. _Il.,_ Ilium; _a._ anterior end; _b._ posterior end; _Ia._ ischium; _Pb.,_ pubis; _T,_ tibia; _F,_ fibula; _As.,_ astragalus; _Ca.,_ calcaneum; I, distal portion of the tarsus; i., ii., iii., iv., metatarsal bones.

Taking all these facts together, it is obvious that the view, which was entertained by Mantell and the probability of which was demonstrated by your own distinguished anatomist, Leidy, while much additional evidence in the same direction has been furnished by Professor Cope, that some of these animals may have walked upon their hind legs as birds do, acquires great weight. In fact, there can be no reasonable doubt that one of the smaller forms of the _Ornithoscelida, Compsognathus,_ the almost entire skeleton of which has been discovered in the Solenhofen slates, was a bipedal animal. The parts of this skeleton are somewhat twisted out of their natural relations, but the accompanying figure gives a just view of the general form of _Compsognathus_ and of the proportions of its limbs; which, in some respects, are more completely bird-like than those of other _Ornithoscelida._

Fig. 7.--Restoration of Compsognathus Longipes

We have had to stretch the definition of the class of birds so as to include birds with teeth and birds with paw-like fore limbs and long tails. There is no evidence that _Compsognathus_ possessed feathers; but, if it did, it would be hard indeed to say whether it should be called a reptilian bird or an avian reptile.

As _Compsognathus_ walked upon its hind legs, it must have made tracks like those of birds. And as the structure of the limbs of several of the gigantic _Ornithoscelida,_ such as _Iguanodon,_ leads to the conclusion that they also may have constantly, or occasionally, assumed the same attitude, a peculiar interest attaches to the fact that, in the Wealden strata of England, there are to be found gigantic footsteps, arranged in order like those of the _Brontozoum,_ and which there can be no reasonable doubt were made by some of the _Ornithoscelida,_ the remains of which are found in the same rocks. And, knowing that reptiles that walked upon their hind legs and shared many of the anatomical characters of birds did once exist, it becomes a very important question whether the tracks in the Trias of Massachusetts, to which I referred some time ago, and which formerly used to be unhesitatingly ascribed to birds, may not all have been made by ornithoscelidan reptiles; and whether, if we could obtain the skeletons of the animals which made these tracks, we should not find in them the actual steps of the evolutional process by which reptiles gave rise to birds.

The evidential value of the facts I have brought forward in this Lecture must be neither over nor under estimated. It is not historical proof of the occurrence of the evolution of birds from reptiles, for we have no safe ground for assuming that true birds had not made their appearance at the commencement of the Mesozoic epoch. It is, in fact, quite possible that all these more or less avi-form reptiles of the Mesozoic epochs are not terms in the series of progression from birds to reptiles at all, but simply the more or less modified descendants of Palaeozoic forms through which that transition was actually effected.

We are not in a position to say that the known _Ornithoscelida_ are intermediate in the order of their appearance on the earth between reptiles and birds. All that can be said is that, if independent evidence of the actual occurrence of evolution is producible, then these intercalary forms remove every difficulty in the way of understanding what the actual steps of the process, in the case of birds, may have been.

That intercalary forms should have existed in ancient times is a necessary consequence of the truth of the hypothesis of evolution; and, hence, the evidence I have laid before you in proof of the existence of such forms, is, so far as it goes, in favour of that hypothesis.

There is another series of extinct reptiles which may be said to be intercalary between reptiles and birds, in so far as they combine some of the characters of both these groups; and which, as they possessed the power of flight, may seem, at first sight, to be nearer representatives of the forms by which the transition from the reptile to the bird was effected, than the _Ornithoscelida._

These are the _Pterosauria,_ or Pterodactyles, the remains of which are met with throughout the series of Mesozoic rocks, from the lias to the chalk, and some of which attained a great size, their wings having a span of eighteen or twenty feet. These animals, in the form and proportions of the head and neck relatively to the body, and in the fact that the ends of the jaws were often, if not always, more or less extensively ensheathed in horny beaks, remind us of birds. Moreover, their bones contained air cavities, rendering them specifically lighter, as is the case in most birds. The breast bone was large and keeled, as in most birds and in bats, and the shoulder girdle is strikingly similar to that of ordinary birds. But, it seems to me, that the special resemblance of pterodactyles to birds ends here, unless I may add the entire absence of teeth which characterises the great pterodactyles _(Pteranodon)_ discovered by Professor Marsh. All other known pterodactyles have teeth lodged in sockets. In the vertebral column and the hind limbs there are no special resemblances to birds, and when we turn to the wings they are found to be constructed on a totally different principle from those of birds.

Fig. 8.--Pterodactylus Spectabilis (Von Meyer).

There are four fingers. These four fingers are large, and three of them, those which answer to the thumb and two following fingers in my hand--are terminated by claws, while the fourth is enormously prolonged and converted into a great jointed style. You see at once, from what I have stated about a bird's wing, that there could be nothing less like a bird's wing than this is. It was concluded by general reasoning that this finger had the office of supporting a web which extended between it and the body. An existing specimen proves that such was really the case, and that the pterodactyles were devoid of feathers, but that the fingers supported a vast web like that of a bat's wing; in fact, there can be no doubt that this ancient reptile flew after the fashion of a bat.

Thus, though the pterodactyle is a reptile which has become modified in such a manner as to enable it to fly, and therefore, as might be expected, presents some points of resemblance to other animals which fly; it has, so to speak, gone off the line which leads directly from reptiles to birds, and has become disqualified for the changes which lead to the characteristic organisation of the latter class. Therefore, viewed in relation to the classes of reptiles and birds, the pterodactyles appear to me to be, in a limited sense, intercalary forms; but they are not even approximately linear, in the sense of exemplifying those modifications of structure through which the passage from the reptile to the bird took place.

III. THE DEMONSTRATIVE EVIDENCE OF EVOLUTION

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 now have 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 palaeontology 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 fulfils 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 those points which are absolutely essential to our inquiry.

Let us turn in the first place to the fore-limb. In most quadrupeds, as in ourselves, the fore-arm contains 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 fore arm, 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 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 gristly 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 its 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 fore-arm bone is composed of the coalesced radius and ulna.