In one of his finest stories, Sur La Pierre Blanche, Anatole France has imagined a group of Roman patricians discussing the future of their Empire. The Christians, who are about to rise to power on their ruin, they dismiss with amiable indifference as one of the little passing eccentricities of the religious life of their time. They have not the dimmest prevision, even as the dream of a possibility, that in a century or two the Empire of Rome will lie in the dust, and the cross will tower above all its cities from York to Jerusalem. If we might for a moment endow the animals of the Mesozoic world with AEsopian wisdom, we could imagine some such discussion taking place between a group of Deinosaur patricians. They would reflect with pride on the unshakable empire of the reptiles, and perhaps glance with disdain at two types of animals which hid in the recesses or fled to the hills of the Jurassic world. And before another era of the earth's story opened, the reptile race would be dethroned, and these hunted and despised and feeble eccentricities of Mesozoic life would become the masters of the globe.
These two types of organisms were the bird and the mammal. Both existed in the Jurassic, and the mammals at least had many representatives in the Triassic. In other words, they existed, with all their higher organisation, during several million years without attaining power. The mammals remained, during at least 3,000,000 years, a small and obscure caste, immensely overshadowed by the small-brained reptiles. The birds, while making more progress, apparently, than the mammals, were far outnumbered by the flying reptiles until the last part of the Mesozoic. Then there was another momentous turn of the wheel of fate, and they emerged from their obscurity to assume the lordship of the globe.
In earlier years, when some serious hesitation was felt by many to accept the new doctrine of evolution, a grave difficulty was found in the circumstance that new types—not merely new species and new genera, but new orders and even sub-classes—appeared in the geological record quite suddenly. Was it not a singular coincidence that in ALL cases the intermediate organisms between one type and another should have wholly escaped preservation? The difficulty was generally due to an imperfect acquaintance with the conditions of the problem. The fossil population of a period is only that fraction of its living population which happened to be buried in a certain kind of deposit under water of a certain depth. We shall read later of insects being preserved in resin (amber), and we have animals (and even bacteria) preserved in trees from the Coal-forests. Generally speaking, however, the earth has buried only a very minute fraction of its land-population. Moreover, only a fraction of the earth's cemeteries have yet been opened. When we further reflect that the new type of organism, when it first appears, is a small and local group, we see what the chances are of our finding specimens of it in a few scattered pages of a very fragmentary record of the earth's life. We shall see that we have discovered only about ten skeletons or fragments of skeletons of the men who lived on the earth before the Neolithic period; a stretch of some hundreds of thousands of years, recorded in the upper strata of the earth.
Whatever serious difficulty there ever was in this scantiness of intermediate types is amply met by the fact that every fresh decade of search in the geological tombs brings some to light. We have seen many instances of this—the seed-bearing ferns and flower-bearing cycads, for example, found in the last decade—and will see others. But one of the most remarkable cases of the kind now claims our attention. The bird was probably evolved in the late Triassic or early Jurassic. It appears in abundance, divided into several genera, in the Chalk period. Luckily, two bird-skeletons have been found in the intermediate period, the Jurassic, and they are of the intermediate type, between the reptile and the bird, which the theory of evolution would suggest. But for the fortunate accident of these two birds being embedded in an ancient Bavarian mud-layer, which happened to be opened, for commercial purposes, in the second half of the nineteenth century, critics of evolution—if there still were any in the world of science—might be repeating to-day that the transition from the reptile to the bird was unthinkable in theory and unproven in fact.
The features of the Archaeopteryx ("primitive bird") have been described so often, and such excellent pictorial restorations of its appearance may now be seen, that we may deal with it briefly. We have in it a most instructive combination of the characters of the bird and the reptile. The feathers alone, the imprint of which is excellently preserved in the fine limestone, would indicate its bird nature, but other anatomical distinctions are clearly seen in it. "There is," says Dr. Woodward, "a typical bird's 'merrythought' between the wings, and the hind leg is exactly that of a perching bird." In other words, it has the shoulder-girdle and four-toed foot, as well as the feathers, of a bird. On the other hand, it has a long tail (instead of a terminal tuft of feathers as in the bird) consisting of twenty-one vertebrae, with the feathers springing in pairs from either side; it has biconcave vertebrae, like the fishes, amphibia, and reptiles; it has teeth in its jaws; and it has three complete fingers, free and clawed, on its front limbs.
As in the living Peripatus, therefore, we have here a very valuable connecting link between two very different types of organisms. It is clear that one of the smaller reptiles—the Archaeopteryx is between a pigeon and a crow in size—of the Triassic period was the ancestor of the birds. Its most conspicuous distinction was that it developed a coat of feathers. A more important difference between the bird and the reptile is that the heart of the bird is completely divided into four chambers, but, as we saw, this probably occurred also in the other flying reptiles. It may be said to be almost a condition of the greater energy of a flying animal. When the heart has four complete chambers, the carbonised blood from the tissues of the body can be conveyed direct to the lungs for purification, and the aerated blood taken direct to the tissues, without any mingling of the two. In the mud-fish and amphibian, we saw, the heart has two chambers (auricles) above, but one (ventricle) below, in which the pure and impure blood mingle. In the reptiles a partition begins to form in the lower chamber. In the turtle it is so nearly complete that the venous and the arterial blood are fairly separated; in the crocodile it is quite complete, though the arteries are imperfectly arranged. Thus the four-chambered heart of the bird and mammal is not a sudden and inexplicable development. Its advantage is enormous in a cold climate. The purer supply of blood increases the combustion in the tissues, and the animal maintains its temperature and vitality when the surrounding air falls in temperature. It ceases to be "cold-blooded."
But the bird secures a further advantage, and here it outstrips the flying reptile. The naked skin of the Pterosaur would allow the heat to escape so freely when the atmosphere cooled that a great strain would be laid on its vitality. A man lessens the demand on his vitality in cold regions by wearing clothing. The bird somehow obtained clothing, in the shape of a coat of feathers, and had more vitality to spare for life-purposes in a falling temperature. The reptile is strictly limited to one region, the bird can pass from region to region as food becomes scarce.
The question of the origin of the feathers can be discussed only from the speculative point of view, as they are fully developed in the Archaeopteryx, and there is no approach toward them in any other living or fossil organism. But a long discussion of the problem has convinced scientific men that the feathers are evolved from the scales of the reptile ancestor. The analogy between the shedding of the coat in a snake and the moulting of a bird is not uninstructive. In both cases the outer skin or epidermis is shedding an old growth, to be replaced by a new one. The covering or horny part of the scale and the feather are alike growths from the epidermis, and the initial stages of the growth have certain analogies. But beyond this general conviction that the feather is a development of the scale, we cannot proceed with any confidence. Nor need we linger in attempting to trace the gradual modification of the skeleton, owing to the material change in habits. The horny beak and the reduction of the toes are features we have already encountered in the reptile, and the modification of the pelvis, breast-bone, and clavicle are a natural outcome of flight.
In the Chalk period we find a large number of bird remains, of about thirty different species, and in some respects they resume the story of the evolution of the bird. They are widely removed from our modern types of birds, and still have teeth in the jaws. They are of two leading types, of which the Ichthyornis and Hesperornis are the standard specimens. The Ichthyornis was a small, tern-like bird with the power of flight strongly developed, as we may gather from the frame of its wings and the keel-shaped structure of its breast-bone. Its legs and feet were small and slender, and its long, slender jaws had about twenty teeth on each side at the bottom. No modern bird has teeth; though the fact that in some modern species we find the teeth appearing in a rudimentary form is another illustration of the law that animals tend to reproduce ancestral features in their development. A more reptilian character in the Ichthyornis group is the fact that, unlike any modern bird, but like their reptile ancestors, they had biconcave vertebrae. The brain was relatively poor. We are still dealing with a type intermediate in some respects between the reptile and the modern bird. The gannets, cormorants, and pelicans are believed to descend from some branch of this group.
The other group of Cretaceous birds, of the Hesperornis type, show an actual degeneration of the power of flight through adaptation to an environment in which it was not needed, as happened, later, in the kiwi of New Zealand, and is happening in the case of the barn-yard fowl. These birds had become divers. Their wings had shrunk into an abortive bone, while their powerful legs had been peculiarly fitted for diving. They stood out at right angles to the body, and seem to have developed paddles. The whole frame suggests that the bird could neither walk nor fly, but was an excellent diver and swimmer. Not infrequently as large as an ostrich (five to six feet high), with teeth set in grooves in its jaws, and the jaws themselves joined as in the snake, with a great capacity of bolting its prey, the Hesperornis would become an important element in the life of the fishes. The wing-fingers have gone, and the tail is much shortened, but the grooved teeth and loosely jointed jaws still point back to a reptilian ancestry.
These are the only remains of bird-life that we find in the Mesozoic rocks. Admirably as they illustrate the evolution of the bird from the reptile, they seem to represent a relatively poor development and spread of one of the most advanced organisms of the time. It must be understood that, as we shall see, the latter part of the Chalk period does not belong to the depression, the age of genial climate, which I call the Middle Ages of the earth, but to the revolutionary period which closes it. We may say that the bird, for all its advances in organisation, remains obscure and unprosperous as long as the Age of Reptiles lasts. It awaits the next massive uplift of the land and lowering of temperature.
In an earlier chapter I hinted that the bird and the mammal may have been the supreme outcomes of the series of disturbances which closed the Primary Epoch and devastated its primitive population. As far as the bird is concerned, this may be doubted on the ground that it first appears in the upper or later Jurassic, and is even then still largely reptilian in character. We must remember, however, that the elevation of the land and the cold climate lasted until the second part of the Triassic, and it is generally agreed that the bird may have been evolved in the Triassic. Its slow progress after that date is not difficult to understand. The advantage of a four-chambered heart and warm coat would be greatly reduced when the climate became warmer. The stimulus to advance would relax. The change from a coat of scales to a coat of feathers obviously means adaptation to a low temperature, and there is nothing to prevent us from locating it in the Triassic, and indeed no later known period of cold in which to place it.
It is much clearer that the mammals were a product of the Permian revolution. They not only abound throughout the Jurassic, in which they are distributed in more than thirty genera, but they may be traced into the Triassic itself. Both in North America and Europe we find the teeth and fragments of the jaws of small animals which are generally recognised as mammals. We cannot, of course, from a few bones deduce that there already, in the Triassic, existed an animal with a fully developed coat of fur and an apparatus, however crude, in the breast for suckling the young. But these bones so closely resemble the bones of the lowest mammals of to-day that this seems highly probable. In the latter part of the long period of cold it seems that some reptile exchanged its scales for tufts of hair, developed a four-chambered heart, and began the practice of nourishing the young from its own blood which would give the mammals so great an ascendancy in a colder world.
Nor can we complain of any lack of evidence connecting the mammal with a reptile ancestor. The earliest remains we find are of such a nature that the highest authorities are still at variance as to whether they should be classed as reptilian or mammalian. A skull and a fore limb from the Triassic of South Africa (Tritylodon and Theriodesmus) are in this predicament. It will be remembered that we divided the primitive reptiles of the Permian period into two great groups, the Diapsids and Synapsids (or Theromorphs). The former group have spread into the great reptiles of the Jurassic; the latter have remained in comparative obscurity. One branch of these Theromorph reptiles approach the mammals so closely in the formation of the teeth that they have received the name "of the Theriodonts", or "beast-toothed" reptiles. Their teeth are, like those of the mammals, divided into incisors, canines (sometimes several inches long), and molars; and the molars have in some cases developed cusps or tubercles. As the earlier remains of mammals which we find are generally teeth and jaws, the resemblance of the two groups leads to some confusion in classifying them, but from our point of view it is not unwelcome. It narrows the supposed gulf between the reptile and the mammal, and suggests very forcibly the particular branch of the reptiles to which we may look for the ancestry of the mammals. We cannot say that these Theriodont reptiles were the ancestors of the mammals. But we may conclude with some confidence that they bring us near to the point of origin, and probably had at least a common ancestor with the mammals.
The distribution of the Theriodonts suggests a further idea of interest in regard to the origin of the mammals. It would be improper to press this view in the present state of our knowledge, yet it offers a plausible theory of the origin of the mammals. The Theriodonts seem to have been generally confined to the southern continent, Gondwana Land (Brazil to Australia), of which an area survives in South Africa. It is there also that we find the early disputed remains of mammals. Now we saw that, during the Permian, Gondwana Land was heavily coated with ice, and it seems natural to suppose that the severe cold which the glacial fields would give to the whole southern continent was the great agency in the evolution of the highest type of the animal world. From this southern land the new-born mammals spread northward and eastward with great rapidity. Fitted as they were to withstand the rigorous conditions which held the reptiles and amphibia in check, they seemed destined to attain at once the domination of the earth. Then, as we saw, the land was revelled once more until its surface broke into a fresh semi-tropical luxuriance, and the Deinosaurs advanced to their triumph. The mammals shrank into a meagre and insignificant population, a scattered tribe of small insect-eating animals, awaiting a fresh refrigeration of the globe.
The remains of these interesting early mammals, restricted, as they generally are, to jaws and teeth and a few other bones that cannot in themselves be too confidently distinguished from those of certain reptiles, may seem insufficient to enable us to form a picture of their living forms. In this, however, we receive a singular and fortunate assistance. Some of them are found living in nature to-day, and their distinctly reptilian features would, even if no fossil remains were in existence, convince us of the evolution of the mammals.
The southern continent on which we suppose the mammals to have originated had its eastern termination in Australia. New Zealand seems to have been detached early in the Mesozoic, and was never reached by the mammals. Tasmania was still part of the Australian continent. To this extreme east of the southern continent the early mammals spread, and then, during either the Jurassic or the Cretaceous, the sea completed its inroad, and severed Australia permanently from the rest of the earth. The obvious result of this was to shelter the primitive life of Australia from invasion by higher types, especially from the great carnivorous mammals which would presently develop. Australia became, in other words, a "protected area," in which primitive types of life were preserved from destruction, and were at the same time sheltered from those stimulating agencies which compelled the rest of the world to advance. "Advance Australia" is the fitting motto of the present human inhabitants of that promising country; but the standard of progress has been set up in a land which had remained during millions of years the Chinese Empire of the living world. Australia is a fragment of the Middle Ages of the earth, a province fenced round by nature at least three million years ago and preserving, amongst its many invaluable types of life, representatives of that primitive mammal population which we are seeking to understand.
It is now well known that the Duckbill or Platypus (Ornithorhyncus) and the Spiny Anteater (Echidna) of Australia and Tasmania—with one representative of the latter in New Guinea, which seems to have been still connected—are semi-reptilian survivors of the first animals to suckle their young. Like the reptiles they lay tough-coated eggs and have a single outlet for the excreta, and they have a reptilian arrangement of the bones of the shoulder-girdle; like the mammals, they have a coat of hair and a four-chambered heart, and they suckle the young. Even in their mammalian features they are, as the careful research of Australian zoologists has shown, of a transitional type. They are warm-blooded, but their temperature is much lower than that of other mammals, and varies appreciably with the temperature of their surroundings. [*] Their apparatus for suckling the young is primitive. There are no teats, and the milk is forced by the mother through simple channels upon the breast, from which it is licked by the young. The Anteater develops her eggs in a pouch. They illustrate a very early stage in the development of a mammal from a reptile; and one is almost tempted to see in their timorous burrowing habits a reminiscence of the impotence of the early mammals after their premature appearance in the Triassic.
The next level of mammal life, the highest level that it attains in Australia (apart from recent invasions), is the Marsupial. The pouched animals (kangaroo, wallaby, etc.) are the princes of pre-human life in Australia, and represent the highest point that life had reached when that continent was cut off from the rest of the world. A few words on the real significance of the pouch, from which they derive their name, will suffice to explain their position in the story of evolution.
Among the reptiles the task of the mother ends, as a rule, with the laying of the egg. One or two modern reptiles hatch the eggs, or show some concern for them, but the characteristic of the reptile is to discharge its eggs upon the warm earth and trouble no further about its young. It is a reminiscence of the warm primitive earth. The bird and mammal, born of the cooling of the earth, exhibit the beginning of that link between mother and offspring which will prove so important an element in the higher and later life of the globe. The bird assists the development of the eggs with the heat of her own body, and feeds the young. The mammal develops the young within the body, and then feeds them at the breast.
But there is a gradual advance in this process. The Duckbill lays its eggs just like the reptile, but provides a warm nest for them at the bottom of its burrow. The Anteater develops a temporary pouch in its body, when it lays an egg, and hatches the egg in it. The Marsupial retains the egg in its womb until the young is advanced in development, then transfers the young to the pouch, and forces milk into its mouth from its breasts. The real reason for this is that the Marsupial falls far short of the higher mammals in the structure of the womb, and cannot fully develop its young therein. It has no placenta, or arrangement by which the blood-vessels of the mother are brought into connection with the blood-vessels of the foetus, in order to supply it with food until it is fully developed. The Marsupial, in fact, only rises above the reptile in hatching the egg within its own body, and then suckling the young at the breast.
These primitive mammals help us to reconstruct the mammal life of the Mesozoic Epoch. The bones that we have are variously described in geological manuals as the remains of Monotremes, Marsupials, and Insectivores. Many of them, if not most, were no doubt insect-eating animals, but there is no ground for supposing that what are technically known as Insectivores (moles and shrews) existed in the Mesozoic. On the other hand, the lower jaw of the Marsupial is characterised by a peculiar hooklike process, and this is commonly found in Mesozoic jaws. This circumstance, and the witness of Australia, permit us, perhaps, to regard the Jurassic mammals as predominantly marsupial. It is more difficult to identify Monotreme remains, but the fact that Monotremes have survived to this day in Australia, and the resemblance of some of the Mesozoic teeth to those found for a time in the young Duckbill justify us in assuming that a part of the Mesozoic mammals correspond to the modern Monotremes. Not single specimen of any higher, or placental, mammal has yet been found in the whole Mesozoic Era.
We must, however, beware of simply transferring to the Mesozoic world the kinds of Monotremes and Marsupials which we know in nature to-day. In some of the excellent "restorations" of Mesozoic life which are found in recent illustrated literature the early mammal is represented with an external appearance like that of the Duckbill. This is an error, as the Duckbill has been greatly modified in its extremities and mouth-parts by its aquatic and burrowing habits. As we have no complete skeletons of these early mammals we must abstain from picturing their external appearance. It is enough that the living Monotreme and Marsupial so finely illustrate the transition from a reptilian to a mammalian form. There may have been types more primitive than the Duckbill, and others between the Duckbill and the Marsupial. It seems clear, at least, that two main branches, the Monotremes and Marsupials, arose from the primitive mammalian root. Whether either of these became in turn the parent of the higher mammals we will inquire later. We must first consider the fresh series of terrestrial disturbances which, like some gigantic sieve, weeded out the grosser types of organisms, and cleared the earth for a rapid and remarkable expansion of these primitive birds and mammals.
We have attended only to a few prominent characters in tracing the line of evolution, but it will be understood that an advance in many organs of the body is implied in these changes. In the lower mammals the diaphragm, or complete partition between the organs of the breast and those of the abdomen, is developed. It is not a sudden and mysterious growth, and its development in the embryo to-day corresponds to the suggestion of its development which the zoologist gathers from the animal series. The ear also is now fully developed. How far the fish has a sense of hearing is not yet fully determined, but the amphibian certainly has an organ for the perception of waves of sound. Parts of the discarded gill-arches are gradually transformed into the three bones of the mammal's internal ear; just as other parts are converted into mouth cartilages, and as—it is believed—one of the gill clefts is converted into the Eustachian tube. In the Monotreme and Marsupial the ear-hole begins to be covered with a shell of cartilage; we have the beginning of the external ear. The jaws, which are first developed in the fish, now articulate more perfectly with the skull. Fat-glands appear in the skin, and it is probably from a group of these that the milk-glands are developed. The origin of the hairs is somewhat obscure. They are not thought to be, like the bird's feathers, modifications of the reptile's scales, but to have been evolved from other structures in the skin, possibly under the protection of the scales.
My purpose is, however, rather to indicate the general causes of the onward advance of life than to study organs in detail—a vast subject—or construct pedigrees. We therefore pass on to consider the next great stride that is taken by the advancing life of the earth. Millions of years of genial climate and rich vegetation have filled the earth with a prolific and enormously varied population. Over this population the hand of natural selection is outstretched, as it were, and we are about to witness another gigantic removal of older types of life and promotion of those which contain the germs of further advance. As we have already explained, natural selection is by no means inactive during these intervening periods of warmth. We have seen the ammonites and reptiles, and even the birds and mammals, evolve into hundreds of species during the Jurassic period. The constant evolution of more effective types of carnivores and their spread into new regions, the continuous changes in the distribution of land and water, the struggle for food in a growing population, and a dozen other causes, are ever at work. But the great and comprehensive changes in the face of the earth which close the eras of the geologist seem to give a deeper and quicker stimulus to its population and result in periods of especially rapid evolution. Such a change now closes the Mesozoic Era, and inaugurates the age of flowering plants, of birds, and of mammals.
In accordance with the view of the later story of the earth which was expressed on an earlier page, we now come to the second of the three great revolutions which have quickened the pulse of life on the earth. Many men of science resent the use of the word revolution, and it is not without some danger. It was once thought that the earth was really shaken at times by vast and sudden cataclysms, which destroyed its entire living population, so that new kingdoms of plants and animals had to be created. But we have interpreted the word revolution in a very different sense. The series of changes and disturbances to which we give the name extended over a period of hundreds of thousands of years, and they were themselves, in some sense, the creators of new types of organisms. Yet they are periods that stand out peculiarly in the comparatively even chronicle of the earth. The Permian period transformed the face of the earth; it lifted the low-lying land into a massive relief, drew mantles of ice over millions of miles of its surface, set volcanoes belching out fire and fumes in many parts, stripped it of its great forests, and slew the overwhelming majority of its animals. On the scale of geological time it may be called a revolution.
It must be confessed that the series of disturbances which close the Secondary and inaugurate the Tertiary Era cannot so conveniently be summed up in a single formula. They begin long before the end of the Mesozoic, and they continue far into the Tertiary, with intervals of ease and tranquillity. There seems to have been no culminating point in the series when the uplifted earth shivered in a mantle of ice and snow. Yet I propose to retain for this period—beginning early in the Cretaceous (Chalk) period and extending into the Tertiary—the name of the Cretaceous Revolution. I drew a fanciful parallel between the three revolutions which have quickened the earth since the sluggish days of the Coal-forest and the three revolutionary movements which have changed the life of modern Europe. It will be remembered that, whereas the first of these European revolutions was a sharp and massive upheaval, the second consisted in a more scattered and irregular series of disturbances, spread over the fourth and fifth decades of the nineteenth century; but they amounted, in effect, to a revolution.
So it is with the Cretaceous Revolution. In effect it corresponds very closely to the Permian Revolution. On the physical side it includes a very considerable rise of the land over the greater part of the globe, and the formation of lofty chains of mountains; on the botanical side it means the reduction of the rich Mesozoic flora to a relatively insignificant population, and the appearance and triumphant spread of the flowering plants, on the zoological side it witnesses the complete extinction of the Ammonites, Deinosaurs, and Pterosaurs, an immense reduction of the reptile world generally, and a victorious expansion of the higher insects, birds, and mammals; on the climatic side it provides the first definite evidence of cold zones of the earth and cold seasons of the year, and seems to represent a long, if irregular, period of comparative cold. Except, to some extent, the last of these points, there is no difference of opinion, and therefore, from the evolutionary point of view, the Cretaceous period merits the title of a revolution. All these things were done before the Tertiary period opened.
Let us first consider the fundamental and physical aspect of this revolution, the upheaval of the land. It began about the close of the Jurassic period. Western and Central Europe emerged considerably from the warm Jurassic sea, which lay on it and had converted it into an archipelago. In North-western America also there was an emergence of large areas of land, and the Sierra and Cascade ranges of mountains were formed about the same time. For reasons which will appear later we must note carefully this rise of land at the very beginning of the Cretaceous period.
However, the sea recovered its lost territory, or compensation for it, and the middle of the Cretaceous period witnessed a very considerable extension of the waters over America, Europe, and southern Asia. The thick familiar beds of chalk, which stretch irregularly from Ireland to the Crimea, and from the south of Sweden to the south of France, plainly tell of an overlying sea. As is well known, the chalk consists mainly of the shells or outer frames of minute one-celled creatures (Thalamophores) which float in the ocean, and form a deep ooze at its bottom with their discarded skeletons. What depth this ocean must have been is disputed, and hardly concerns us. It is clear that it must have taken an enormous period for microscopic shells to form the thick masses of chalk which cover so much of southern and eastern England. On the lowest estimates the Cretaceous period, which includes the deposit of other strata besides chalk, lasted about three million years. And as people like to have some idea of the time since these things happened, I may add that, on the lowest estimate (which most geologists would at least double), it is about three million years since the last stretches of the chalk-ocean disappeared from the surface of Europe.
But while our chalk cliffs conjure up a vision of England lying deep—at least twenty or thirty fathoms deep—below a warm ocean, in which gigantic Ammonites and Belemnites and sharks ply their deadly trade, they also remind us of the last phase of the remarkable life of the earth's Middle Ages. In the latter part of the Cretaceous the land rises. The chalk ocean of Europe is gradually reduced to a series of inland seas, separated by masses and ridges of land, and finally to a series of lakes of brackish water. The masses of the Pyrenees and Alps begin to rise; though it will not be until a much later date that they reach anything like their present elevation. In America the change is even greater. A vast ridge rises along the whole western front of the continent, lifting and draining it, from Alaska to Cape Horn. It is the beginning of the Rocky Mountains and the Andes. Even during the Cretaceous period there had been rich forests of Mesozoic vegetation covering about a hundred thousand square miles in the Rocky Mountains region. Europe and America now begin to show their modern contours.
It is important to notice that this great uprise of the land and the series of disturbances it entails differ from those which we summed up in the phrase Permian Revolution. The differences may help us to understand some of the changes in the living population. The chief difference is that the disturbances are more local, and not nearly simultaneous. There is a considerable emergence of land at the end of the Jurassic, then a fresh expansion of the sea, then a great rise of mountains at the end of the Cretaceous, and so on. We shall find our great mountain-masses (the Pyrenees, Alps, Himalaya, etc.) rising at intervals throughout the whole of the Tertiary Era. However, it suffices for the moment to observe that in the latter part of the Mesozoic and early part of the Tertiary there were considerable upheavals of the land in various regions, and that the Mesozoic Era closed with a very much larger proportion of dry land, and a much higher relief of the land, than there had been during the Jurassic period. The series of disturbances was, says Professor Chamberlin, "greater than any that had occurred since the close of the Palaeozoic."
From the previous effect of the Permian upheaval, and from the fact that the living population is now similarly annihilated or reduced, we should at once expect to find a fresh change in the climate of the earth. Here, however, our procedure is not so easy. In the Permian age we had solid proof in the shape of vast glaciated regions. It is claimed by continental geologists that certain early Tertiary beds in Bavaria actually prove a similar, but smaller, glaciation in Europe, but this is disputed. Other beds may yet be found, but we saw that there was not a general upheaval, as there had been in the Permian, and it is quite possible that there were few or no ice-fields. We do not, in fact, know the causes of the Permian icefields. We are thrown upon the plant and animal remains, and seem to be in some danger of inferring a cold climate from the organic remains, and then explaining the new types of organisms by the cold climate. This, of course, we shall not do. The difficulty is made greater by the extreme disinclination of many recent geologists, and some recent botanists who have too easily followed the geologists, to admit a plain climatic interpretation of the facts. Let us first see what the facts are.
In the latter part of the Jurassic we find three different zones of Ammonites: one in the latitude of the Mediterranean, one in the latitude of Central Europe, and one further north. Most geologists conclude that these differences indicate zones of climate (not hitherto indicated), but it cannot be proved, and we may leave the matter open. At the same time the warm-loving corals disappear from Europe, with occasional advances. It is said that they are driven out by the disturbance of the waters, and, although this would hardly explain why they did not spread again in the tranquil chalk-ocean, we may again leave the point open.
In the early part of the Cretaceous, however, the Angiosperms (flowering plants) suddenly break into the chronicle of the earth, and spread with great rapidity. They appear abruptly in the east of the North American continent, in the region of Virginia and Maryland. They are small in stature and primitive in structure. Some are of generalised forms that are now unknown; some have leaves approaching those of the oak, willow, elm, maple, and walnut; some may be definitely described as fig, sassafras, aralia, myrica, etc. Eastern America, it may be recalled, is much higher than western until the close of the Cretaceous period. The Angiosperms do not spread much westward; they appear next in Greenland, and, before the middle of the Cretaceous, in Portugal. They have travelled over the North Atlantic continent, or what remains of it. The process seems very rapid as we write it, but it must be remembered that the first half of the Cretaceous period means a million or a million and a half years.
The cycads, and even the conifers, shrink before the higher type of tree. The landscape, in Europe and America, begins to wear a modern aspect. Long before the end of the Cretaceous most of the modern genera of Angiosperm trees have developed. To the fig and sassafras are now added the birch, beech, oak, poplar, walnut, willow, ivy, mulberry, holly, laurel, myrtle, maple, oleander, magnolia, plane, bread-fruit, and sweet-gum. Most of the American trees of to-day are known. The sequoias (the giant Californian trees) still represent the conifers in great abundance, with the eucalyptus and other plants that are now found only much further south. The ginkgoes struggle on for a time. The cycads dwindle enormously. Of 700 specimens in one early Cretaceous deposit only 96 are Angiosperms; of 460 species in a later deposit about 400 are Angiosperms. They oust the cycads in Europe and America, as the cycads and conifers had ousted the Cryptogams. The change in the face of the earth would be remarkable. Instead of the groves of palm-like cycads, with their large and flower-like fructifications, above which the pines and firs and cypresses reared their sombre forms, there were now forests of delicate-leaved maples, beeches, and oaks, bearing nutritious fruit for the coming race of animals. Grasses also and palms begin in the Cretaceous; though the grasses would at first be coarse and isolated tufts. Even flowers, of the lily family (apparently), are still detected in the crushed and petrified remains.
We will give some consideration later to the evolution of the Angiosperms. For the moment it is chiefly important to notice a feature of them to which the botanist pays less attention. In his technical view the Angiosperm is distinguished by the structure of its reproductive apparatus, its flowers, and some recent botanists wonder whether the key to this expansion of the flowering plants may not be found in a development of the insect world and of its relation to vegetation. In point of fact, we have no geological indication of any great development of the insects until the Tertiary Era, when we shall find them deploying into a vast army and producing their highest types. In any case, such a view leaves wholly unexplained the feature of the Angiosperms which chiefly concerns us. This is that most of them shed the whole of their leaves periodically, as the winter approaches. No such trees had yet been known on the earth. All trees hitherto had been evergreen, and we need a specific and adequate explanation why the earth is now covered, in the northern region, with forests of trees which show naked boughs and branches during a part of the year.
The majority of palaeontologists conclude at once, and quite confidently, from this rise and spread of the deciduous trees, that a winter season has at length set in on the earth, and that this new type of vegetation appears in response to an appreciable lowering of the climate. The facts, however, are somewhat complex, and we must proceed with caution. It would seem that any general lowering of the temperature of the earth ought to betray itself first in Greenland, but the flora of Greenland remains far "warmer," so to say, than the flora of Central Europe is to-day. Even toward the close of the Cretaceous its plants are much the same as those of America or of Central Europe. Its fossil remains of that time include forty species of ferns, as well as cycads, ginkgoes, figs, bamboos, and magnolias. Sir A. Geikie ventures to say that it must then have enjoyed a climate like that of the Cape or of Australia to-day. Professor Chamberlin finds its flora like that of "warm temperate" regions, and says that plants which then flourished in latitude 72 degrees are not now found above latitude 30 degrees.
There are, however, various reasons to believe that it is unsafe to draw deductions from the climate of Greenland. There is, it is true, some exaggeration in the statement that its climate was equivalent to that of Central Europe. The palms which flourished in Central Europe did not reach Greenland, and there are differences in the northern Molluscs and Echinoderms which—like the absence of corals above the north of England—point to a diversity of temperature. But we have no right to expect that there would be the same difference in temperature between Greenland and Central Europe as we find to-day. If the warm current which is now diverted to Europe across the Atlantic—the Gulf Stream—had then continued up the coast of America, and flowed along the coast of the land that united America and Europe, the climatic conditions would be very different from what they are. There is a more substantial reason. We saw that during the Mesozoic the Arctic continent was very largely submerged, and, while Europe and America rise again at the end of the Cretaceous, we find no rise of the land further north. A difference of elevation would, in such a world, make a great difference in temperature and moisture.
Let us examine the animal record, however, before we come to any conclusion. The chronicle of the later Cretaceous is a story of devastation. The reduction of the cyeads is insignificant beside the reduction or annihilation of the great animals of the Mesozoic world. The skeletons of the Deinosaurs become fewer and fewer as we ascend the upper Cretaceous strata. In the uppermost layer (Laramie) we find traces of a last curious expansion—the group of horned reptiles, of the Triceratops type, which we described as the last of the great reptiles. The Ichthyosaurs and Plesiosaurs vanish from the waters. The "sea-serpents" (Mososaurs) pass away without a survivor. The flying dragons, large and small, become entirely extinct. Only crocodiles, lizards, turtle, and snakes cross the threshold of the Tertiary Era. In one single region of America (Puerco beds) some of the great reptiles seem to be making a last stand against the advancing enemy in the dawn of the Tertiary Era, but the exact date of the beds is disputed, and in any case their fight is soon over. Something has slain the most formidable race that the earth had yet known, in spite of its marvellous adaptation to different environments in its innumerable branches.
We turn to the seas, and find an equal carnage among some of its most advanced inhabitants. The great cuttlefish-like Belemnites and the whole race of the Ammonites, large and small, are banished from the earth. The fall of the Ammonites is particularly interesting, and has inspired much more or less fantastic speculation. The shells begin to assume such strange forms that observers speak occasionally of the "convulsions" or "death-contortions" of the expiring race. Some of the coiled shells take on a spiral form, like that of a snail's shell. Some uncoil the shell, and seem to be returning toward the primitive type. A rich eccentricity of frills and ornamentation is found more or less throughout the whole race. But every device—if we may so regard these changes—is useless, and the devastating agency of the Cretaceous, whatever it was, removes the Ammonites and Belemnites from the scene. The Mollusc world, like the world of plants and of reptiles, approaches its modern aspect.
In the fish world, too, there is an effective selection in the course of the Cretaceous. All the fishes of modern times, except the large family of the sharks, rays, skates, and dog-fishes (Elasmobranchs), the sturgeon and chimaera, the mud-fishes, and a very few other types, are Teleosts, or bony-framed fishes—the others having cartilaginous frames. None of the Teleosts had appeared until the end of the Jurassic. They now, like the flowering plants on land, not only herald the new age, but rapidly oust the other fishes, except the unconquerable shark. They gradually approach the familiar types of Teleosts, so that we may say that before the end of the Cretaceous the waters swarmed with primitive and patriarchal cod, salmon, herring, perch, pike, bream, eels, and other fishes. Some of them grew to an enormous size. The Portheus, an American pike, seems to have been about eight feet long; and the activity of an eight-foot pike may be left to the angler's imagination. All, however, are, as evolution demands, of a generalised and unfamiliar type: the material out of which our fishes will be evolved.
Of the insects we have very little trace in the Cretaceous. We shall find them developing with great richness in the following period, but, imperfect as the record is, we may venture to say that they were checked in the Cretaceous. There were good conditions for preserving them, but few are preserved. And of the other groups of invertebrates we need only say that they show a steady advance toward modern types. The sea-lily fills the rocks no longer; the sea-urchin is very abundant. The Molluscs gain on the more lowly organised Brachiopods.
To complete the picture we must add that higher types probably arose in the later Cretaceous which do not appear in the records. This is particularly true of the birds and mammals. We find them spreading so early in the Tertiary that we must put back the beginning of the expansion to the Cretaceous. As yet, however, the only mammal remains we find are such jaws and teeth of primitive mammals as we have already described. The birds we described (after the Archaeopteryx) also belong to the Cretaceous, and they form another of the doomed races. Probably the modern birds were already developing among the new vegetation on the higher ground.
These are the facts of Cretaceous life, as far as the record has yielded them, and it remains for us to understand them. Clearly there has been a great selective process analogous to, if not equal to, the winnowing process at the end of the Palaeozoic. As there has been a similar, if less considerable, upheaval of the land, we are at once tempted to think that the great selective agency was a lowering of the temperature. When we further find that the most important change in the animal world is the destruction of the cold-blooded reptiles, which have no concern for the young, and the luxuriant spread of the warm-blooded animals, which do care for their young, the idea is greatly confirmed. When we add that the powerful Molluscs which are slain, while the humbler Molluscs survive, are those which—to judge from the nautilus and octopus—love warm seas, the impression is further confirmed. And when we finally reflect that the most distinctive phenomenon of the period is the rapid spread of deciduous trees, it would seem that there is only one possible interpretation of the Cretaceous Revolution.
This interpretation—that cold was the selecting agency—is a familiar idea in geological literature, but, as I said, there are recent writers who profess reserve in regard to it, and it is proper to glance at, or at least look for, the alternatives.
Before doing so let us be quite clear that here we have nothing to do with theories of the origin of the earth. The Permian cold—which, however, is universally admitted—is more or less entangled in that controversy; the Cretaceous cold has no connection with it. Whatever excess of carbon-dioxide there may have been in the early atmosphere was cleared by the Coal-forests. We must set aside all these theories in explaining the present facts.
It is also useful to note that the fact that there have been great changes in the climate of the earth in past time is beyond dispute. There is no denying the fact that the climate of the earth was warm from the Arctic to the Antarctic in the Devonian and Carboniferous periods: that it fell considerably in the Permian: that it again became at least "warm temperate" (Chamberlin) from the Arctic to the Antarctic in the Jurassic, and again in the Eocene: that some millions of square miles of Europe and North America were covered with ice and snow in the Pleistocene, so that the reindeer wandered where palms had previously flourished and the vine flourishes to-day; and that the pronounced zones of climate which we find today have no counterpart in any earlier age. In view of these great and admitted fluctuations of the earth's temperature one does not see any reason for hesitating to admit a fall of temperature in the Cretaceous, if the facts point to it.
On the other hand, the alternative suggestions are not very convincing. We have noticed one of these suggestions in connection with the origin of the Angiosperms. It hints that this may be related to developments of the insect world. Most probably the development of the characteristic flowers of the Angiosperms is connected with an increasing relation to insects, but what we want to understand especially is the deciduous character of their leaves. Many of the Angiosperms are evergreen, so that it cannot be said that the one change entailed the other. In fact, a careful study of the leaves preserved in the rocks seems to show the deciduous Angiosperms gaining on the evergreens at the end of the Cretaceous. The most natural, it not the only, interpretation of this is that the temperature is falling. Deciduous trees shed their leaves so as to check their transpiration when a season comes on in which they cannot absorb the normal amount of moisture. This may occur either at the on-coming of a hot, dry season or of a cold season (in which the roots absorb less). Everything suggests that the deciduous tree evolved to meet an increase of cold, not of heat.
Another suggestion is that animals and plants were not "climatically differentiated" until the Cretaceous period; that is to say, that they were adapted to all climates before that time, and then began to be sensitive to differences of climate, and live in different latitudes. But how and why they should suddenly become differentiated in this way is so mysterious that one prefers to think that, as the animal remains also suggest, there were no appreciable zones of climate until the Cretaceous. The magnolia, for instance, flourished in Greenland in the early Tertiary, and has to live very far south of it to-day. It is much simpler to assume that Greenland changed—as a vast amount of evidence indicates—than that the magnolia changed.
Finally, to explain the disappearance of the Mesozoic reptiles without a fall in temperature, it is suggested that they were exterminated by the advancing mammals. It is assumed that the spreading world of the Angiospermous plants somewhere met the spread of the advancing mammals, and opened out a rich new granary to them. This led to so powerful a development of the mammals that they succeeded in overthrowing the reptiles.
There are several serious difficulties in the way of this theory. The first and most decisive is that the great reptiles have practically disappeared before the mammals come on the scene. Only in one series of beds (Puerco) in America, representing an early period of the Tertiary Era, do we find any association of their remains; and even there it is not clear that they were contemporary. Over the earth generally the geological record shows the great reptiles dying from some invisible scourge long before any mammal capable of doing them any harm appears; even if we suppose that the mammal mainly attacked the eggs and the young. We may very well believe that more powerful mammals than the primitive Mesozoic specimens were already developed in some part of the earth—say, Africa—and that the rise of the land gave them a bridge across the Mediterranean to Europe. Probably this happened; but the important point is that the reptiles were already almost extinct. The difficulty is even greater when we reflect that it is precisely the most powerful reptiles (Deinosaurs) and least accessible reptiles (Pterosaurs, Ichthyosaurs, etc.) which disappear, while the smaller land and water reptiles survive and retreat southward—where the mammals are just as numerous. That assuredly is not the effect of an invasion of carnivores, even if we could overlook the absence of such carnivores from the record until after the extinction of the reptiles in most places.
I have entered somewhat fully into this point, partly because of its great interest, but partly lest it be thought that I am merely reproducing a tradition of geological literature without giving due attention to the criticisms of recent writers. The plain and common interpretation of the Cretaceous revolution—that a fall in temperature was its chief devastating agency—is the only one that brings harmony into all the facts. The one comprehensive enemy of that vast reptile population was cold. It was fatal to the adult because he had a three-chambered heart and no warm coat; it was fatal to the Mesozoic vegetation on which, directly or indirectly, he fed; it was fatal to his eggs and young because the mother did not brood over the one or care for the other. It was fatal to the Pterosaurs, even if they were warm-blooded, because they had no warm coats and did not (presumably) hatch their eggs; and it was equally fatal to the viviparous Ichthyosaurs. It is the one common fate that could slay all classes. When we find that the surviving reptiles retreat southward, only lingering in Europe during the renewed warmth of the Eocene and Miocene periods, this interpretation is sufficiently confirmed. And when we recollect that these things coincide with the extinction of the Ammonites and Belemnites, and the driving of their descendants further south, as well as the rise and triumph of deciduous trees, it is difficult to see any ground for hesitating.
But we need not, and must not, imagine a period of cold as severe, prolonged, and general as that of the Permian period. The warmth of the Jurassic period is generally attributed to the low relief of the land, and the very large proportion of water-surface. The effect of this would be to increase the moisture in the atmosphere. Whether this was assisted by any abnormal proportion of carbon-dioxide, as in the Carboniferous, we cannot confidently say. Professor Chamberlin observes that, since the absorbing rock-surface was greatly reduced in the Jurassic, the carbon-dioxide would tend to accumulate in its atmosphere, and help to explain the high temperature. But the great spread of vegetation and the rise of land in the later Jurassic and the Cretaceous would reduce this density of the atmosphere, and help to lower the temperature.
It is clear that the cold would at first be local. In fact, it must be carefully realised that, when we speak of the Jurassic period as a time of uniform warmth, we mean uniform at the same altitude. Everybody knows the effect of rising from the warm, moist sea-level to the top of even a small inland elevation. There would be such cooler regions throughout the Jurassic, and we saw that there were considerable upheavals of land towards its close. To these elevated lands we may look for the development of the Angiosperms, the birds, and the mammals. When the more massive rise of land came at the end of the Cretaceous, the temperature would fall over larger areas, and connecting ridges would be established between one area and another. The Mesozoic plants and animals would succumb to this advancing cold. What precise degree of cold was necessary to kill the reptiles and Cephalopods, yet allow certain of the more delicate flowering plants to live, is yet to be determined. The vast majority of the new plants, with their winter sleep, would thrive in the cooler air, and, occupying the ground of the retreating cycads and ginkgoes would prepare a rich harvest for the coming birds and mammals.