AIIMS: Evolution Time Series: Tertiary
The Tertiary Era
We have already traversed nearly nine-tenths of the story of terrestrial life, without counting the long and obscure Archaean period, and still find ourselves in a strange and unfamiliar earth. With the close of the Chalk period, however, we take a long stride in the direction of the modern world. The Tertiary Era will, in the main, prove a fresh period of genial warmth and fertile low-lying regions. During its course our deciduous trees and grasses will mingle with the palms and pines over the land, our flowers will begin to brighten the landscape, and the forms of our familiar birds and mammals, even the form of man, will be discernible in the crowds of animals. At its close another mighty period of selection will clear the stage for its modern actors.
A curious reflection is prompted in connection with this division of the earth's story into periods of relative prosperity and quiescence, separated by periods of disturbance. There was--on the most modest estimate--a stretch of some fifteen million years between the Cambrian and the Permian upheavals. On the same chronological scale the interval between the Permian and Cretaceous revolutions was only about seven million years, and the Tertiary Era will comprise only about three million years. One wonders if the Fourth (Quaternary) Era in which we live will be similarly shortened. Further, whereas the earth returned after each of the earlier upheavals to what seems to have been its primitive condition of equable and warm climate, it has now entirely departed from that condition, and exhibits very different zones of climate and a succession of seasons in the year. One wonders what the climate of the earth will become long before the expiration of those ten million years which are usually assigned as the minimum period during which the globe will remain habitable.
It is premature to glance at the future, when we are still some millions of years from the present, but it will be useful to look more closely at the facts which inspire this reflection. From what we have seen, and shall further see, it is clear that, in spite of all the recent controversy about climate among our geologists, there has undeniably been a progressive refrigeration of the globe. Every geologist, indeed, admits “oscillations of climate,” as Professor Chamberlin puts it. But amidst all these oscillations we trace a steady lowering of the temperature. Unless we put a strained and somewhat arbitrary interpretation on the facts of the geological record, earlier ages knew nothing of our division of the year into pronounced seasons and of the globe into very different climatic zones. It might plausibly be suggested that we are still living in the last days of the Ice-Age, and that the earth may be slowly returning to a warmer condition. Shackleton, it might be observed, found that there has been a considerable shrinkage of the south polar ice within the period of exploration. But we shall find that a difference of climate, as compared with earlier ages, was already evident in the middle of the Tertiary Era, and it is far more noticeable to-day.
We do not know the causes of this climatic evolution--the point will be considered more closely in connection with the last Ice-Age--but we see that it throws a flood of light on the evolution of organisms. It is one of the chief incarnations of natural selection. Changes in the distribution of land and water and in the nature of the land-surface, the coming of powerful carnivores, and other agencies which we have seen, have had their share in the onward impulsion of life, but the most drastic agency seems to have been the supervention of cold. The higher types of both animals and plants appear plainly in response to a lowering of temperature. This is the chief advantage of studying the story of evolution in strict connection with the geological record. We shall find that the record will continue to throw light on our path to the end, but, as we are now about to approach the most important era of evolution, and as we have now seen so much of the concrete story of evolution, it will be interesting to examine briefly some other ways of conceiving that story.
We need not return to the consideration of the leading schools of evolution, as described in a former chapter. Nothing that we have seen will enable us to choose between the Lamarckian and the Weismannist hypothesis; and I doubt if anything we are yet to see will prove more decisive. The dispute is somewhat academic, and not vital to a conception of evolution. We shall, for instance, presently follow the evolution of the horse, and see four of its toes shrink and disappear, while the fifth toe is enormously strengthened. In the facts themselves there is nothing whatever to decide whether this evolution took place on the lines suggested by Weismann, or on the lines suggested by Lamarck and accepted by Darwin. It will be enough for us merely to establish the fact that the one-toed horse is an evolved descendant of a primitive five-toed mammal, through the adaptation of its foot to running on firm ground, its teeth and neck to feeding on grasses, and so on:
On the other hand, the facts we have already seen seem to justify the attitude of compromise I adopted in regard to the Mutationist theory. It would be an advantage in many ways if we could believe that new species arose by sudden and large variations (mutations) of the young from the parental type. In the case of many organs and habits it is extremely difficult to see how a gradual development, by a slow accentuation of small variations, is possible. When we further find that experimenters on living species can bring about such mutations, and when we reflect that there must have been acute disturbances in the surroundings of animals and plants sometimes, we are disposed to think that many a new species may have arisen in this way. On the other hand, while the palaeontological record can never prove that a species arose by mutations, it does sometimes show that species arise by very gradual modification. The Chalk period, which we have just traversed, affords a very clear instance. One of our chief investigators of the English Chalk, Dr. Rowe, paid particular attention to the sea-urchins it contains, as they serve well to identify different levels of chalk. He discovered, not merely that they vary from level to level, but that in at least one genus (Micraster) he could trace the organism very gradually passing from one species to another, without any leap or abruptness. It is certainly significant that we find such cases as this precisely where the conditions of preservation are exceptionally good. We must conclude that species arise, probably, both by mutations and small variations, and that it is impossible to say which class of species has been the more numerous.
Conceptions of Evolution
There remain one or two conceptions of evolution which we have not hitherto noticed, as it was advisable to see the facts first. One of these is the view--chiefly represented in this country by Professor Henslow--that natural selection has had no part in the creation of species; that the only two factors are the environment and the organism which responds to its changes. This is true enough in the sense that, as we saw, natural selection is not an action of nature on the “fit,” but on the unfit or less fit. But this does not in the least lessen the importance of natural selection. If there were not in nature this body of destructive agencies, to which we apply the name natural selection, there would be little--we cannot say no--evolution. But the rising carnivores, the falls of temperature, etc. that we have studied, have had so real, if indirect, an influence on the development of life that we need not dwell on this.
Another school, or several schools, while admitting the action of natural selection, maintain that earlier evolutionists have made nature much too red in tooth and claw. Dr. Russel Wallace from one motive, and Prince Krapotkin from another, have insisted that the triumphs of war have been exaggerated, and the triumphs of peace, or of social co-operation, far too little appreciated. It will be found that such writers usually base their theory on life as we find it in nature to-day, where the social principle is highly developed in many groups of animals. This is most misleading, since social co-operation among animals, as an instrument of progress, is (geologically speaking) quite a recent phenomenon. Nearly every group of animals in which it is found belongs, to put it moderately, to the last tenth of the story of life, and in some of the chief instances the animals have only gradually developed social life. * The first nine-tenths of the chronicle of evolution contain no indication of social life, except--curiously enough--in such groups as the Sponges, Corals, and Bryozoa, which are amongst the least progressive in nature. We have seen plainly that during the overwhelmingly greater part of the story of life the predominant agencies of evolution were struggle against adverse conditions and devouring carnivores; and we shall find them the predominant agencies throughout the
Thus the social nature of man is sometimes quoted as one of the chief causes of his development. It is true that it has much to do with his later development, but we shall see that the statement that man was from the start a social being is not at all warranted by the facts. On the other hand, it may be pointed out that the ants and termites had appeared in the Mesozoic. We shall see some evidence that the remarkable division of labour which now characterises their life did not begin until a much later period, so that we have no evidence of social life in the early stages.
Yet we must protest against the exaggerated estimate of the conscious pain which so many read into these millions of years of struggle. Probably there was no consciousness at all during the greater part of the time. The wriggling of the worm on which you have accidentally trodden is no proof whatever that you have caused conscious pain. The nervous system of an animal has been so evolved as to respond with great disturbance of its tissue to any dangerous or injurious assault. It is the selection of a certain means of self-preservation. But at what level of life the animal becomes conscious of this disturbance, and “feels pain,” it is very difficult to determine. The subject is too vast to be opened here. In a special investigation of it * I concluded that there is no proof of the presence of any degree of consciousness in the invertebrate world even in the higher insects; that there is probably only a dull, blurred, imperfect consciousness below the level of the higher mammals and birds; and that even the consciousness of an ape is something very different from what educated Europeans, on the ground of their own experience, call consciousness. It is too often forgotten that pain is in proportion to consciousness. We must beware of such fallacies as transferring our experience of pain to a Mesozoic reptile, with an ounce or two of cerebrum to twenty tons of muscle and bone.
One other view of evolution, which we find in some recent and reputable works (such as Professor Geddes and Thomson's “Evolution,” 1911), calls for consideration. In the ordinary Darwinian view the variations of the young from their parents are indefinite, and spread in all directions. They may continue to occur for ages without any of them proving an advantage to their possessors. Then the environment may change, and a certain variation may prove an advantage, and be continuously and increasingly selected. Thus these indefinite variations may be so controlled by the environment during millions of years that the fish at last becomes an elephant or a man. The alternative view, urged by a few writers, is that the variations were “definitely directed.” The phrase seems merely to complicate the story of evolution with a fresh and superfluous mystery. The nature and precise action of this “definite direction” within the organism are quite unintelligible, and the facts seem explainable just as well--or not less imperfectly--without as with this mystic agency. Radiolaria, Sponges, Corals, Sharks, Mudfishes, Duckbills, etc. do not change (except within the limits of their family) during millions of years, because they keep to an environment to which they are fitted. On the other hand, certain fishes, reptiles, etc. remain in a changing environment, and they must change with it. The process has its obscurities, but we make them darker, it seems to me, with these semi-metaphysical phrases.
Rise of a new Era
It has seemed advisable to take this further glance at the general principles and current theories of evolution before we extend our own procedure into the Tertiary Era. The highest types of animals and plants are now about to appear on the stage of the earth; the theatre itself is about to take on a modern complexion. The Middle Ages are over; the new age is breaking upon the planet. We will, as before, first survey the Tertiary Era as a whole, with the momentous changes it introduces, and then examine, in separate chapters, the more important phases of its life.
It opens, like the preceding and the following era, with “the area of land large and its relief pronounced.” This is the outcome of the Cretaceous revolution. Southern Europe and Southern Asia have risen, and shaken the last masses of the Chalk ocean from their faces; the whole western fringe of America has similarly emerged from the sea that had flooded it. In many parts, as in England (at that time a part of the Continent), there is so great a gap between the latest Cretaceous and the earliest Tertiary strata that these newly elevated lands must evidently have stood out of the waters for a prolonged period. On their cooler plains the tragedy of the extinction of the great reptiles comes to an end. The cyeads and ginkgoes have shrunk into thin survivors of the luxuriant Mesozoic groves. The oak and beech and other deciduous trees spread slowly over the successive lands, amid the glare and thunder of the numerous volcanoes which the disturbance of the crust has brought into play. New forms of birds fly from tree to tree, or linger by the waters; and strange patriarchal types of mammals begin to move among the bones of the stricken reptiles.
But the seas and the rains and rivers are acting with renewed vigour on the elevated lands, and the Eocene period closes in a fresh age of levelling. Let us put the work of a million years or so in a sentence. The southern sea, which has been confined almost to the limits of our Mediterranean by the Cretaceous upheaval, gradually enlarges once more. It floods the north-west of Africa almost as far as the equator; it covers most of Italy, Turkey, Austria, and Southern Russia; it spreads over Asia Minor, Persia, and Southern Asia, until it joins the Pacific; and it sends a long arm across the Franco-British region, and up the great valley which is now the German Ocean.
From earlier chapters we now expect to find a warmer climate, and the record gives abundant proof of it. To this period belongs the “London Clay,” in whose thick and--to the unskilled eye--insignificant bed the geologist reads the remarkable story of what London was two or three million years ago. It tells us that a sea, some 500 or 600 feet deep, then lay over that part of England, and fragments of the life of the period are preserved in its deposit. The sea lay at the mouth of a sub-tropical river on whose banks grew palms, figs, ginkgoes, eucalyptuses, almonds, and magnolias, with the more familiar oaks and pines and laurels. Sword-fishes and monstrous sharks lived in the sea. Large turtles and crocodiles and enormous “sea-serpents” lingered in this last spell of warmth that Central Europe would experience. A primitive whale appeared in the seas, and strange large tapir-like mammals--remote ancestors of our horses and more familiar beasts--wandered heavily on the land. Gigantic primitive birds, sometimes ten feet high, waded by the shore. Deposits of the period at Bournemouth and in the Isle of Wight tell the same story of a land that bore figs, vines, palms, araucarias, and aralias, and waters that sheltered turtles and crocodiles. The Parisian region presented the same features.
Traces of Southern Sea
In fact, one of the most characteristic traces of the southern sea which then stretched from England to Africa in the south and India in the east indicates a warm climate. It will be remembered that the Cretaceous ocean over Southern Europe had swarmed with the animalcules whose dead skeletons largely compose our chalk-beds. In the new southern ocean another branch of these Thalamophores, the Nummulites, spreads with such portentous abundance that its shells--sometimes alone, generally with other material--make beds of solid limestone several thousand feet in thickness. The pyramids are built of this nummulitic limestone. The one-celled animal in its shell is, however, no longer a microscopic grain. It sometimes forms wonderful shells, an inch or more in diameter, in which as many as a thousand chambers succeed each other, in spiral order, from the centre. The beds containing it are found from the Pyrenees to Japan.
That this vast warm ocean, stretching southward over a large part of what is now the Sahara, should give a semitropical aspect even to Central Europe and Asia is not surprising. But this genial climate was still very general over the earth. Evergreens which now need the warmth of Italy or the Riviera then flourished in Lapland and Spitzbergen. The flora of Greenland--a flora that includes magnolias, figs, and bamboos--shows us that its temperature in the Eocene period must have been about 30 degrees higher than it is to-day. * The temperature of the cool Tyrol of modern Europe is calculated to have then been between 74 and 81 degrees F. Palms, cactuses, aloes, gum-trees, cinnamon trees, etc. flourished in the latitude of Northern France. The forests that covered parts of Switzerland which are now buried in snow during a great part of the year were like the forests one finds in parts of India and Australia to-day. The climate of North America, and of the land which still connected it with Europe, was correspondingly genial.
This indulgent period (the Oligocene, or later part of the Eocene), scattering a rich and nutritious vegetation with great profusion over the land, led to a notable expansion of animal life. Insects, birds, and mammals spread into vast and varied groups in every land. Had any of the great Mesozoic reptiles survived, the warmer age might have enabled them to dispute the sovereignty of the advancing mammals. But nothing more formidable than the turtle, the snake, and the crocodile (confined to the waters) had crossed the threshold of the Tertiary Era, and the mammals and birds had the full advantage of the new golden age. The fruits of the new trees, the grasses which now covered the plains, and the insects which multiplied with the flowers afforded a magnificent diet. The herbivorous mammals became a populous world, branching into numerous different types according to their different environments. The horse, the elephant, the camel, the pig, the deer, the rhinoceros gradually emerge out of the chaos of evolving forms. Behind them, hastening the course of their evolution, improving their speed, arms, and armour, is the inevitable carnivore. He, too, in the abundance of food, grows into a vast population, and branches out toward familiar types. We will devote a chapter presently to this remarkable phase of the story of evolution.
Periods of Tertiary Era
The Tertiary Era is divided by geologists into four periods: The Eocene, Oligocene, Miocene, and Pliocene. “Cene” is our barbaric way of expressing the Greek word for “new,” and the classification is meant to mark the increase of new (or modern and actual) types of life in the course of the Tertiary Era. Many geologists, however, distrust the classification, and are disposed to divide the Tertiary into two periods. From our point of view, at least, it is advisable to do this. The first and longer half of the Tertiary is the period in which the temperature rises until Central Europe enjoys the climate of South Africa; the second half is the period in which the land gradually rises, and the temperature falls, until glaciers and sheets of ice cover regions where the palm and fig had flourished.
The rise of the land had begun in the first half of the Tertiary, but had been suspended. The Pyrenees and Apennines had begun to rise at the end of the Eocene, straining the crust until it spluttered with volcanoes, casting the nummulitic sea off large areas of Southern Europe. The Nummulites become smaller and less abundant. There is also some upheaval in North America, and a bridge of land begins to connect the north and south, and permit an effective mingling of their populations. But the advance is, as I said, suspended, and the Oligocene period maintains the golden age. With the Miocene period the land resumes its rise.
A chill is felt along the American coast, showing a fall in the temperature of the Atlantic. In Europe there is a similar chill, and a more obvious reason for it. There is an ascending movement of the whole series of mountains from Morocco and the Pyrenees, through the Alps, the Caucasus, and the Carpathians, to India and China. Large lakes still lie over Western Europe, but nearly the whole of it emerges from the ocean. The Mediterranean still sends an arm up France, and with another arm encircles the Alpine mass; but the upheaval continues, and the great nummulitic sea is reduced to a series of extensive lakes, cut off both from the Atlantic and Pacific. The climate of Southern Europe is probably still as genial as that of the Canaries to-day. Palms still linger in the landscape in reduced numbers.
The last part of the Tertiary, the Pliocene, opens with a slight return of the sea. The upheaval is once more suspended, and the waters are eating into the land. There is some foundering of land at the south-western tip of Europe; the “Straits of Gibraltar” begin to connect the Mediterranean with the Atlantic, and the Balearic Islands, Corsica, and Sardinia remain as the mountain summits of a submerged land. Then the upheaval is resumed, in nearly every part of the earth.