Having established the facts, he proceeded to enquire into the theory. There was now a new method for investigating such problems, the method of embryology, which, practically, had not been available to Oken, and of which neither Cuvier nor Owen had made proper use. By putting together the investigations of a number of embryologists, by adding to these himself, and, lastly, by interpreting the facts which his investigations into comparative anatomy had brought to light, he shewed that the vertebral theory could not be maintained. He shewed, by these methods, that, though both skull and vertebral column are segmented, the one and the other, after an early stage, are fashioned on lines so different as to exclude the possibility of regarding the details of each as mere modifications of a common type. "The spinal column and the skull start from the same primitive condition, whence they immediately begin to diverge." "It may be true to say that there is a primitive identity of structure between the spinal or vertebral column and the skull; but it is no more true that the adult skull is a modified vertebral column than it would be to affirm that the vertebral column is a modified skull." Taking the embryological facts, he shewed that the skull arose out of elements quite different from those of the vertebral column. The notochord alone is common to both. The skull is built up of longitudinal cartilaginous pieces, now known as the "parachordals" and "trabeculæ," of sense capsules enclosing the nose and ear, and of various roofing bones. In the historical development of the skull three grades become apparent; a primitive stage, as seen in Amphioxus, where there is nothing but a fibrous investment of the nervous structures; a cartilaginous grade, as seen in the skate or shark, where the skull is formed of cartilage, very imperfectly hardened by earthy deposits; a bony stage, seen in most of the higher animals. He shewed that in actual development of the higher animals these historical grades are repeated, the skull being at first a mere membranous or fibrous investment of the developing nervous masses, then becoming cartilaginous, and, lastly, bony. He made some important prophetic remarks as to the probable importance that future embryological work would give to the distinction between cartilage and membrane bones—a prophecy that has been more than fully realised by the investigations of Hertwig and of others. Our present knowledge of the skull differs from Huxley's conception practically only in a fuller knowledge of details. We know now that throughout the series there is a primitive set of structures common to all animals higher in the scale than Amphioxus, and forming the base and lateral walls of the skull. This is termed the Chondrocraninm, because it is laid down in cartilage; it is composed of the separate elements which Huxley indicated, and, in different animals, as Huxley suggested, the exact limits of the ossification of the primitive cartilages differ in extent, but occur in homologous situations. This primitive skull is roofed over by a series of membrane bones which have no connection in origin with the other portions of the skull, and which have no representative in the vertebral column, but which are the direct descendants of the bony scales clothing the external skin in cartilaginous fishes. In one respect only was Huxley erroneous. Partly by inadvertence, and partly because the minute details of vertebrate embryology became really familiar to zoölogists only after the elaborate work of Balfour of Cambridge, Huxley, in his account of the formation of the first beginnings of the skeleton in the embryo, made confusion between the walls of the primitive groove, which, in reality, give rise to the nervous structures, and those embryonic tissues which form the skeletal system.

The next great piece of work which we may take as typical of Huxley's contributions to vertebrate anatomy, is his classical study on the classification of birds. The great group of birds contains a larger number of species than is known in any other group of vertebrates, and, in this vast assemblage of forms there is strikingly little anatomical difference. The ostrich and the humming-bird might perhaps be taken as types of the extremest differences to be found, and yet, although these differ in size, plumage, adaptations, habits, mode of life, and almost everything that can separate living things, the two conform so closely to the common type of bird structure that knowledge of the anatomy of one would be a sufficient guide, down to minute details, for dissection of the other. None the less, there are hundreds of thousands of species of birds between these two types. It is not surprising that to reduce this vast assemblage of similar creatures to an ordered system of classification has proved one of the most difficult tasks attempted by zoölogists. Before Huxley, it had been attempted by a number of distinguished zoölogists; but, for the most part, these had relied too much on merely external characters and on superficial modifications in obvious relation to habits. When Huxley, in the course of a set of lectures on Comparative Anatomy, was about to approach the subject of birds he was asked by a zoölogist how he proposed to treat them. "I intend," he replied, "to treat them as extinct animals." By that he meant that it was his purpose to make a prolonged study of their skeletal structures the basis of his grouping, following the lines which Cuvier, Owen, and he himself had pursued so successfully in the case of the fossil remains of vertebrates. The result was that this first systematic study of even one set of the anatomical characters of the group completely reformed the method by which all subsequent workers have tried to grapple with the problem; ornithology was raised from a process akin to stamp-collecting to a reasoned scientific study. The immediate practical results were equally important. He was able to shew that among the innumerable known forms there were three grades of structure. The lowest had already been recognised and named by Haeckel; it consisted of the Saururæ, or reptile-like, birds, and contained a single fossil form, Archæopteryx, distinguished from all living birds by the presence of a hand-like wing in which the metacarpal bones were well developed and freely movable, and by the possession of a long lizard-like tail actually exceeding in length the remainder of the spinal column. The next group of Ratites, although it contained only the Ostrich, Rhea, Emu, Cassowary, and Apteryx, he shewed to be equivalent in anatomical coherence to the third great group of Carinates, which includes the vast majority of living birds. In his arrangement of the latter group, he laid most stress on the characters of the bony structures which form the palate, and by this simple means was able to lay down clearly at least the main lines of a natural classification of the group.

Huxley's work upon birds, like his work in many other branches of anatomy, has been so overlaid by the investigations of subsequent zoölogists that it is easy to overlook its importance. His employment of the skeleton as the basis of classification was succeeded by the work of others who made a similar use of the muscular anatomy, of the intestinal canal, of the windpipe, of the tendons of the feet, and many other structures which display anatomical modifications in different birds. The modern student finds that all these new sets of facts are much greater in bulk than the work of Huxley, and it is easy for him to remain in ignorance that they were all suggested and inspired by the method which Huxley employed. He finds that further research has supplanted some of Huxley's conclusions, and it is easy for him to remain in ignorance that the conclusions themselves suggested the investigations which have modified them. Huxley's anatomical work was essentially living and stimulating, and too often it has become lost to sight simply because of the vast superstructures of new facts to which it gave rise.

Closely associated with vertebrate anatomy is the subject of geographical distribution. In 1857 the study of this important department of zoölogy was placed on a scientific basis, practically for the first time, by a memoir on the geographical distribution of birds published in the Journal of the Linnæan Society of London. It was known in a general way that different kinds of creatures were found in different parts of the world, but little attempt had been made to map out the world into regions characterised by their animal and vegetable inhabitants, as the political divisions of the world are characterised by their different governments and policies. Mr. Sclater, who two years later became secretary of the Zoölogical Society of London, in his memoir introduced the subject in the following words:

"It is a well-known and universally acknowledged fact that we can choose two portions of the globe of which the respective fauna and flora shall be so different that we should not be far wrong in supposing them to have been the result of distinct creations. Assuming, then, that there are, or may be, more areas of creation than one, the question naturally arises how many of them are there, and what are their respective extents and boundaries; or, in other words, what are the most natural primary ontological divisions of the earth's surface?"

Mr. Sclater's answer was that there are six great regions; Neotropical, Nearctic, Palæarctic, Ethiopian, Indian, and Australian, and his answer, with minor alterations and the addition of a great wealth of detail, has been accepted by zoölogy.

Two years later, however, Darwin gave a new meaning and a new importance to Sclater's work, by the new interpretation he caused to be placed on the words "centres of creation." Sclater's facts and areas remained the same; Darwin rejected the idea of separate creations in the older sense of the words, and laid stress on the impossibility of accounting for the resemblances within a region and for the differences between regions by climatic differences and so forth. He raised the questions of modes of dispersal and of barriers to dispersal, of similarities due to common descent, and of the modifying results produced by isolation. He gave, in fact, a theory of the "creations" which Mr. Sclater had shewn to be a probable assumption. It was in the nature of things that Huxley should make a contribution to a set of problems so novel and of so much importance to zoölogy. In 1868, in the course of a memoir on the anatomy of the gallinaceous birds and their allies, he made a useful attempt, nearly the first of its kind, to correlate anatomical facts with geographical distribution. Having shewn the diverging lines of anatomical structure that existed in the group of creatures he had been considering, he went on to shew that there was a definite relation between the varieties of structure and the different positions on the surface of the globe occupied at the present time by the creatures in question. He made, in fact, the geographical position a necessary part of the whole idea of a species or of a group, and so introduced a conception which has become a permanent part of zoölogical science.

With regard to the number and limits of the zoölogical regions into which the world may be divided, Huxley raised a number of problems which have not yet reached a full solution. Mr. Sclater had divided the world into six great regions: the Nearctic, including the continent of North America, with an overlap into what is called South America by geographers; the Palæarctic, comprising Europe and the greater part of Asia; the Oriental, containing certain southern portions of Asia, such as India south of the Himalayas and many of the adjacent islands; the Ethiopian, including Africa, except north of the Sahara, and Madagascar; the Australian, containing Australia and New Zealand and some of the more southeastern of the islands of Malay; the Neotropical, including South America. Huxley first called attention to certain noteworthy resemblances between the Neotropical and the Australian regions of Sclater, and held that a primary division of the world was into Arctogæa, comprising the great land masses of the Northern Hemisphere with a part of their extension across the equator, and Notogæa, which contained Australia but not New Zealand and South America. Although this acute suggestion has not been generally accepted as a modification of Mr. Sclater's scheme, it called attention in a striking fashion to some very remarkable features in the distribution of animals. Subsequent writers have considerably extended Huxley's conception of the similarities to be found among the more southern land areas. They have pointed out that the most striking idea of the distribution of land and water on the surface of the globe is to be got by considering the globe alternately from one pole and from the other. In the south, a clump of ice-bound land, well within the Antarctic Circle, surrounds the pole. All else is a wide domain of ocean broken only where tapering and isolated tongues of land, South America, the Cape, Australia, lean down from the great land masses of the north. On the other hand, all the great land masses expand in the Northern Hemisphere, and shoulder one another round the North Pole. America is separated from Asia only by the shallowest and narrowest of straits; an elevation of a few fathoms would unite Greenland with Europe. Science points definitely to some part of the great northern land area as the centre of life for at least the larger terrestrial forms of life. We know that these arose successively, primitive birds like the ostriches being older than higher forms like the parrots and singing birds; the pouched marsupials preceding the antelopes and the lion; the lemurs coming before the man-like apes. Each wave of life spread over the whole area producing after its kind; then, pressing round the northern land area, it met a thousand different conditions of environment, different foods, enemies, and climates, and broke up into different genera and species. But there was never a wave of life that was not followed by another wave. In the struggle for existence between the newer and the older forms, the older forms were gradually driven southwards towards the diverging fringes of the land masses. The vanquished left behind them on the field of battle only their bones, to become fossils. Sometimes succeeding waves swept along to the extreme limits of the land, and many early types were utterly destroyed. But others found sanctuary in the ends of the South, and such survivors of older and earlier types of life cause a similarity between the southern lands that Huxley called Notogæa, although the extent of his region must be increased.

Recently, however, there has been a recurrence to Huxley's suggested union of South America and Australia, based on new evidence of a direct kind, quite different from that which had just been given. Various groups of naturalists have stated that there are similarities between the invertebrate inhabitants of Australia and of South America of a kind which makes the existence of a direct land connection in the Southern Hemisphere extremely probable. Moreover, Ameghino has recently described some marsupial fossils from South America which, he states, belong to the Australian group of Dasyuridæ, and Oldfield Thomas has described a new mammal from South America which is unlike the opossums of America and like the diprotodonts of Australia. So that, while the general opinion has been against Huxley's division, Notogæa, in the strict meaning which he gave to it, there has recently been an opinion growing in its favour.

Huxley also made minor alterations in Mr. Sclater's scheme by forming an additional circumpolar region for the Northern Hemisphere, and by elevating New Zealand into a separate region, distinct from Australia. On these points there is a balance of opinion against his views.