Lectures on Evolution / Essay #3 from "Science and Hebrew Tradition"

by Thomas Henry Huxley

But I think it may have an especial interest if, instead of dealing with these examples, which would require a great deal of tedious osteological detail, I take the case of birds and reptiles; groups which, at the present day, are so clearly distinguished from one another that there are perhaps no classes of animals which, in popular apprehension, are more completely separated. Existing birds, as you are aware, are covered with feathers; their anterior extremities, specially and peculiarly modified, are converted into wings by the aid of which most of them are able to fly; they walk upright upon two legs; and these limbs, when they are considered anatomically, present a great number of exceedingly remarkable peculiarities, to which I may have occasion to advert incidentally as I go on, and which are not met with, even approximately, in any existing forms of reptiles. On the other hand, existing reptiles have no feathers. They may have naked skins, or be covered with horny scales, or bony plates, or with both. They possess no wings; they neither fly by means of their fore-limbs, nor habitually walk upright upon their hind-limbs; and the bones of their legs present no such modifications as we find in birds. It is impossible to imagine any two groups more definitely and distinctly separated, notwithstanding certain characters which they possess in common.

As we trace the history of birds back in time, we find their remains, sometimes in great abundance, throughout the whole extent of the tertiary rocks; but, so far as our present knowledge goes, the birds of the tertiary rocks retain the same essential characters as the birds of the present day. In other words, the tertiary birds come within the definition of the class constituted by existing birds, and are as much separated from reptiles as existing birds are. Not very long ago no remains of birds had been found below the tertiary rocks, and I am not sure but that some persons were prepared to demonstrate that they could not have existed at an earlier period. But, in the course of the last few years, such remains have been discovered in England; though, unfortunately, in so imperfect and fragmentary a condition, that it is impossible to say whether they differed from existing birds in any essential character or not. In your country the development of the cretaceous series of rocks is enormous; the conditions under which the later cretaceous strata have been deposited are highly favourable to the preservation of organic remains; and the researches, full of labour and risk, which have been carried on by Professor Marsh in these cretaceous rocks of Western America, have rewarded him with the discovery of forms of birds of which we had hitherto no conception. By his kindness, I am enabled to place before you a restoration of one of these extraordinary birds, every part of which can be thoroughly justified by the more or less complete skeletons, in a very perfect state of preservation, which he has discovered. This Hesperornis (Fig. 3), which measured between five and six feet in length, is astonishingly like our existing divers or grebes in a great many respects; so like them indeed that, had the skeleton of Hesperornis been found in a museum without its skull, it probably would have been placed in the same group of birds as the divers and grebes of the present day. [1] But Hesperornis differs from all existing birds, and so far resembles reptiles, in one important particular—it is provided with teeth. The long jaws are armed with teeth which have curved crowns and thick roots (Fig. 4), and are not set in distinct sockets, but are lodged in a groove. In possessing true teeth, the Hesperornis differs from every existing bird, and from every bird yet discovered in the tertiary formations, the tooth-like serrations of the jaws in the Odontopteryx of the London clay being mere processes of the bony substance of the jaws, and not teeth in the proper sense of the word. In view of the characteristics of this bird we are therefore obliged to modify the definitions of the classes of birds and reptiles. Before the discovery of Hesperornis, the definition of the class Aves based upon our knowledge of existing birds might have been extended to all birds; it might have been said that the absence of teeth was characteristic of the class of birds; but the discovery of an animal which, in every part of its skeleton, closely agrees with existing birds, and yet possesses teeth, shows that there were ancient birds which, in respect of possessing teeth, approached reptiles more nearly than any existing bird does, and, to that extent, diminishes the hiatus between the two classes.

Fig. 3—Hesperornis regalis (Marsh)
Fig. 4—Hesperornis regalis (Marsh)
(Side and upper views of half the lower jaw; side and end views
of a vertebra and a separate tooth.)

The same formation has yielded another bird, Ichthyornis (Fig. 5), which also possesses teeth; but the teeth are situated in distinct sockets, while those of Hesperornis are not so lodged. The latter also has such very small, almost rudimentary wings, that it must have been chiefly a swimmer and a diver like a Penguin; while Ichthyornis has strong wings and no doubt possessed corresponding powers of flight. Ichthyornis also differed in the fact that its vertebrae have not the peculiar characters of the vertebrae of existing and of all known tertiary birds, but were concave at each end. This discovery leads us to make a further modification in the definition of the group of birds, and to part with another of the characters by which almost all existing birds are distinguished from reptiles.

Figure. 5—Ichthyornis Dispar (Marsh). Side and upper views
of half the lower jaw; and side and end views of a
vertebra.

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.