With the beginning of life on land we open a new and more important volume of the story of life, and we may take the opportunity to make clearer certain principles or processes of development which we may seem hitherto to have taken for granted. The evolutionary work is too often a mere superficial description of the strange and advancing classes of plants and animals which cross the stage of geology. Why they change and advance is not explained. I have endeavoured to supply this explanation by putting the successive populations of the earth in their respective environments, and showing the continuous and stimulating effect on them of changes in those environments. We have thus learned to decipher some lines of the decalogue of living nature. “Thou shalt have a thick armour,” “Thou shalt be speedy,” “Thou shalt shelter from the more powerful,” are some of the laws of primeval life. The appearance of each higher and more destructive type enforces them with more severity; and in their observance animals branch outward and upward into myriads of temporary or permanent forms.

But there is no consciousness of law and no idea of evading danger. There is not even some mysterious instinct “telling” the animal, as it used to be said, to do certain things. It is, in fact, not strictly accurate to say that a certain change in the environment stimulates animals to advance. Generally speaking, it does not act on the advancing at all, but on the non-advancing, which it exterminates. The procedure is simple, tangible, and unconscious. Two invading arms of the sea meet and pour together their different waters and populations. The habits, the foods, and the enemies of many types of animals are changed; the less fit for the new environment die first, the more fit survive longest and breed most of the new generation. It is so with men when they migrate to a more exacting environment, whether a dangerous trade or a foreign clime. Again, take the case of the introduction of a giant Cephalopod or fish amongst a population of Molluscs and Crustacea. The toughest, the speediest, the most alert, the most retiring, or the least conspicuous, will be the most apt to survive and breed. In hundreds or thousands of generations there will be an enormous improvement in the armour, the speed, the sensitiveness, the hiding practices, and the protective colours, of the animals which are devoured. The “natural selection of the fittest” really means the “natural destruction of the less fit.”

The only point assumed in this is that the young of an animal or plant tend to differ from each other and from their parents. Darwin was content to take this as a fact of common observation, as it obviously is, but later science has thrown some light on the causes of these variations. In the first place, the germs in the parent’s body may themselves be subject to struggle and natural selection, and not share equally in the food-supply. Then, in the case of the higher animals (or the majority of animals), there is a clear source of variation in the fact that the mature germ is formed of certain elements from two different parents, four grandparents, and so on. In the case of the lower animals the germs and larvae float independently in the water, and are exposed to many influences. Modern embryologists have found, by experiment, that an alteration of the temperature or the chemical considerable effect on eggs and larvae. Some recent experiments have shown that such changes may even affect the eggs in the mother’s ovary. These discoveries are very important and suggestive, because the geological changes which we are studying are especially apt to bring about changes of temperature and changes in the freshness or saltiness of water.

Evolution is, therefore, not a “mere description” of the procession of living things; it is to a great extent an explanation of the procession. When, however, we come to apply these general principles to certain aspects of the advance in organisation we find fundamental differences of opinion among biologists, which must be noted. As Sir E. Ray Lankester recently said, it is not at all true that Darwinism is questioned in zoology to-day. It is true only that Darwin was not omniscient or infallible, and some of his opinions are disputed.

Let me introduce the subject with a particular instance of evolution, the flat-fish. This animal has been fitted to survive the terrible struggle in the seas by acquiring such a form that it can lie almost unseen upon the floor of the ocean. The eye on the under side of the body would thus be useless, but a glance at a sole or plaice in a fishmonger’s shop will show that this eye has worked upward to the top of the head. Was the eye shifted by the effort and straining of the fish, inherited and increased slightly in each generation? Is the explanation rather that those fishes in each generation survived and bred which happened from birth to have a slight variation in that direction, though they did not inherit the effect of the parent’s effort to strain the eye? Or ought we to regard this change of structure as brought about by a few abrupt and considerable variations on the part of the young? There you have the three great schools which divide modern evolutionists: Lamarckism, Weismannism, and Mendelism (or Mutationism). All are Darwinians. No one doubts that the flat-fish was evolved from an ordinary fish ­the flat-fish is an ordinary fish in its youth ­or that natural selection (enemies) killed off the old and transitional types and overlooked (and so favoured) the new. It will be seen that the language used in this volume is not the particular language of any one of these schools. This is partly because I wish to leave seriously controverted questions open, and partly from a feeling of compromise, which I may explain.

First, the plain issue between the Mendelians and the other two schools ­whether the passage from species to species is brought about by a series of small variations during a long period or by a few large variations (or “mutations”) in a short period ­is open to an obvious compromise. It is quite possible that both views are correct, in different cases, and quite impossible to find the proportion of each class of cases. We shall see later that in certain instances where the conditions of preservation were good we can sometimes trace a perfectly gradual advance from species to species. Several shellfish have been traced in this way, and a sea-urchin in the chalk has been followed, quite gradually, from one end of a genus to the other. It is significant that the advance of research is multiplying these cases. There is no reason why we may not assume most of the changes of species we have yet seen to have occurred in this way. In fact, in some of the lower branches of the animal world (Radiolaria, Sponges, etc.) there is often no sharp division of species at all, but a gradual series of living varieties.

On the other hand we know many instances of very considerable sudden changes. The cases quoted by Mendelists generally belong to the plant world, but instances are not unknown in the animal world. A shrimp (Artemia) was made to undergo considerable modification, by altering the proportion of salt in the water in which it was kept. Butterflies have been made to produce young quite different from their normal young by subjecting them to abnormal temperature, electric currents, and so on; and, as I said, the most remarkable effects have been produced on eggs and embryos by altering the chemical and physical conditions. Rats ­I was informed by the engineer in charge of the refrigerating room on an Australian liner ­very quickly became adapted to the freezing temperature by developing long hair. All that we have seen of the past changes in the environment of animals makes it probable that these larger variations often occur. I would conclude, therefore, that evolution has proceeded continuously (though by no means universally) through the ages, but there were at times periods of more acute change with correspondingly larger changes in the animal and plant worlds.

In regard to the issue between the Lamarckians and Weismannists ­whether changes acquired by the parent are inherited by the young ­recent experiments again suggest something of a compromise. Weismann says that the body of the parent is but the case containing the germ-plasm, so that all modifications of the living parent body perish with it, and do not affect the germ, which builds the next generation. Certainly, when we reflect that the 70,000 ova in the human mother’s ovary seem to have been all formed in the first year of her life, it is difficult to see how modifications of her muscles or nerves can affect them. Thus we cannot hope to learn anything, either way, by cutting off the tails of cows, and experiments of that kind. But it is acknowledged that certain diseases in the blood, which nourishes the germs, may affect them, and recent experimenters have found that they can reach and affect the germs in the body by other agencies, and so produce inherited modifications in the parent. If this claim is sustained and enlarged, it may be concluded that the greater changes of environment which we find in the geological chronicle may have had a considerable influence of this kind.

See a paper read by Professor Bourne to the Zoological Section of the British Association, 1910. It must be understood that when I speak of Weismannism I do not refer to this whole theory of heredity, which, he acknowledges, has few supporters. The Lamarckian view is represented in Britain by Sir W. Turner and Professor Darwin. In other countries it has a larger proportion of distinguished supporters. On the whole subject see Professor J. A. Thomson’s “Heredity” (1909), Dewar and Finn’s “Making of Species” (1909 ­a Mendelian work), and, for essays by the leaders of each school, “Darwinism and Modern Science” (1909).

The general issue, however, must remain open. The Lamarckian and Weismannist theories are rival interpretations of past events, and we shall not find it necessary to press either. When the fish comes to live on land, for instance, it develops a bony limb out of its fin. The Lamarckian says that the throwing of the weight of the body on the main stem of the fin strengthens it, as practice strengthens the boxer’s arm, and the effect is inherited and increased in each generation, until at last the useless paddle of the fin dies away and the main stem has become a stout, bony column. Weismann says that the individual modification, by use in walking, is not inherited, but those young are favoured which have at birth a variation in the strength of the stem of the fin. As each of these interpretations is, and must remain, purely theoretical, we will be content to tell the facts in such cases. But these brief remarks will enable the reader to understand in what precise sense the facts we record are open to controversy.

Let us return to the chronicle of the earth. We had reached the Devonian age, when large continents, with great inland seas, existed in North America, north-west Europe, and north Asia, probably connected by a continent across the North Atlantic and the Arctic region. South America and South Africa were emerging, and a continent was preparing to stretch from Brazil, through South Africa and the Antarctic, to Australia and India. The expanse of land was, with many oscillations, gaining on the water, and there was much emigration to it from the over-populated seas. When the fish went on land in the Devonian, it must have found a diet (insects, etc.) there, and the insects must have been preceded by a plant population. We have first, therefore, to consider the evolution of the plant, and see how it increases in form and number until it covers the earth with the luxuriant forests of the Carboniferous period.

The plant world, we saw, starts, like the animal world, with a great kingdom of one-celled microscopic representatives, and the same principles of development, to a great extent, shape it into a large variety of forms. Armour-plating has a widespread influence among them. The graceful Diatom is a morsel of plasm enclosed in a flinty box, often with a very pretty arrangement of the pores and markings. The Desmid has a coat of cellulose, and a less graceful coat of cellulose encloses the Peridinean. Many of these minute plants develop locomotion and a degree of sensitiveness (Diatoms, Peridinea, Euglena, etc.). Some (Bacteria) adopt animal diet, and rise in power of movement and sensitiveness until it is impossible to make any satisfactory distinction between them and animals. Then the social principle enters. First we have loose associations of one-celled plants in a common bed, then closer clusters or many-celled bodies. In some cases (Volvox) the cluster, or the compound plant, is round and moves briskly in the water, closely resembling an animal. In most cases, the cells are connected in chains, and we begin to see the vague outline of the larger plant.

When we had reached this stage in the development of animal life, we found great difficulty in imagining how the chief lines of the higher Invertebrates took their rise from the Archaean chaos of early many-celled forms. We have an even greater difficulty here, as plant remains are not preserved at all until the Devonian period. We can only conclude, from the later facts, that these primitive many-celled plants branched out in several different directions. One section (at a quite unknown date) adopted an organic diet, and became the Fungi; and a later co-operation, or life-partnership, between a Fungus and a one-celled Alga led to the Lichens. Others remained at the Alga-level, and grew in great thickets along the sea bottoms, no doubt rivalling or surpassing the giant sea-weeds, sometimes 400 feet long, off the American coast to-day. Other lines which start from the level of the primitive many-celled Algae develop into the Mosses (Bryophyta), Ferns (Pteridophyta), Horsetails (Equisetalia), and Club-mosses (Lycopodiales). The mosses, the lowest group, are not preserved in the rocks; from the other three classes will come the great forests of the Carboniferous period.

The early record of plant-life is so poor that it is useless to speculate when the plant first left the water. We have somewhat obscure and disputed traces of ferns in the Ordovician, and, as they and the Horsetails and Club-mosses are well developed in the Devonian, we may assume that some of the sea-weeds had become adapted to life on land, and evolved into the early forms of the ferns, at least in the Cambrian period. From that time they begin to weave a mantle of sombre green over the exposed land, and to play a most important part in the economy of nature.

We saw that at the beginning of the Devonian there was a considerable rise of the land both in America and Europe, but especially in Europe. A distant spectator at that time would have observed the rise of a chain of mountains in Scotland and a general emergence of land north-western Europe. A continent stretched from Ireland to Scandinavia and North Russia, while most of the rest of Europe, except large areas of Russia, France, Germany, and Turkey, was under the sea. Where we now find our Alps and Pyrénées towering up to the snow-line there were then level stretches of ocean. Even the north-western continent was scooped into great inland seas or lagoons, which stretched from Ireland to Scandinavia, and, as we saw, fostered the development of the fishes.

As the Devonian period progressed the sea gained on the land, and must have restricted the growth of vegetation, but as the lake deposits now preserve the remains of the plants which grow down to their shores, or are washed into them, we are enabled to restore the complexion of the landscape. Ferns, generally of a primitive and generalised character, abound, and include the ferns such as we find in warm countries to-day. Horsetails and Club-mosses already grow into forest-trees. There are even seed-bearing ferns, which give promise of the higher plants to come, but as yet nothing approaching our flower and fruit-bearing trees has appeared. There is as yet no certain indication of the presence of Conifers. It is a sombre and monotonous vegetation, unlike any to be found in any climate to-day.

We will look more closely into its nature presently. First let us see how these primitive types of plants come to form the immense forests which are recorded in our coal-beds. Dr. Russel Wallace has lately represented these forests, which have, we shall see, had a most important influence on the development of life, as somewhat mysterious in their origin. If, however, we again consult the geologist as to the changes which were taking place in the distribution of land and water, we find a quite natural explanation. Indeed, there are now distinguished geologists (e.g. Professor Chamberlin) who doubt if the Coal-forests were so exceptionally luxuriant as is generally believed. They think that the vegetation may not have been more dense than in some other ages, but that there may have been exceptionally good conditions for preserving the dead trees. We shall see that there were; but, on the whole, it seems probable that during some hundreds of thousands of years remarkably dense forests covered enormous stretches of the earth’s surface, from the Arctic to the Antarctic.

The Devonian period had opened with a rise of the land, but the sea eat steadily into it once more, and, with some inconsiderable oscillations of the land, regained its territory. The latter part of the Devonian and earlier part of the Carboniferous were remarkable for their great expanses of shallow water and low-lying land. Except the recent chain of hills in Scotland we know of no mountains. Professor Chamberlin calculates that 20,000,000, or 30,000,000 square miles of the present continental surface of Europe and America were covered with a shallow sea. In the deeper and clearer of these waters the earliest Carboniferous rocks, of limestone, were deposited. The “millstone grit,” which succeeds the “limestone,” indicates shallower water, which is being rapidly filled up with the debris of the land. In a word, all the indications suggest the early and middle Carboniferous as an age of vast swamps, of enormous stretches of land just above or below the sea-level, and changing repeatedly from one to the other. Further, the climate was at the time ­we will consider the general question of climate later ­moist and warm all over the earth, on account of the great proportion of sea-surface and the absence of high land (not to speak of more disputable causes).

These were ideal conditions for the primitive vegetation, and it spread over the swamps with great vigour. To say that the Coal-forests were masses of Ferns, Horsetails, and Club-mosses is a lifeless and misleading expression. The Club-mosses, or Lycopodiales, were massive trees, rising sometimes to a height of 120 feet, and probably averaging about fifty feet in height and one or two feet in diameter. The largest and most abundant of them, the Sigillaria, sent up a scarred and fluted trunk to a height of seventy or a hundred feet, without a branch, and was crowned with a bunch of its long, tapering leaves. The Lépidodendron, its fellow monarch of the forest, branched at the summit, and terminated in clusters of its stiff, needle-like leaves, six’ or seven inches long, like enormous exaggerations of the little cones at the ends of our Club-mosses to-day. The Horsetails, which linger in their dwarfed descendants by our streams to-day, and at their exceptional best (in a part of South America) form slender stems about thirty feet high, were then forest-trees, four to six feet in circumference and sometimes ninety feet in height. These Calamités probably rose in dense thickets from the borders of the lakes, their stumpy leaves spreading in whorls at every joint in their hollow stems. Another extinct tree, the Cordaites, rivalled the Horsetails and Club-mosses in height, and its showers of long and extraordinary leaves, six feet long and six inches in width, pointed to the higher plant world that was to come. Between these gaunt towering trunks the graceful tree-ferns spread their canopies at heights of twenty, forty, and even sixty feet from the ground, and at the base was a dense undergrowth of ferns and fern-like seed-plants. Mosses may have carpeted the moist ground, but nothing in the nature of grass or flowers had yet appeared.

Imagine this dense assemblage of dull, flowerless trees pervaded by a hot, dank atmosphere, with no change of seasons, with no movement but the flying of large and primitive insects among the trees and the stirring of the ferns below by some passing giant salamander, with no song of bird and no single streak of white or red or blue drawn across the changeless sombre green, and you have some idea of the character of the forests that are compressed into our seams of coal. Imagine these forests spread from Spitzbergen to Australia and even, according to the south polar expeditions, to the Antarctic, and from the United States to Europe, to Siberia, and to China, and prolonged during some hundreds of thousands of years, and you begin to realise that the Carboniferous period prepared the land for the coming dynasties of animals. Let some vast and terrible devastation fall upon this luxuriant world, entombing the great multitude of its imperfect forms and selecting the higher types for freer life, and the earth will pass into a new age.

But before we describe the animal inhabitants of these forests, the part that the forests play in the story of life, and the great cataclysm which selects the higher types from the myriads of forms which the warm womb of the earth has poured out, we must at least glance at the evolutionary position of the Carboniferous plants themselves. Do they point downward to lower forms, and upward to higher forms, as the theory of evolution requires? A close inquiry into this would lead us deep into the problems of the modern botanist, but we may borrow from him a few of the results of the great labour he has expended on the subject within the last decade.

Just as the animal world is primarily divided into Invertebrates and Vertebrates, the plant world is primarily divided into a lower kingdom of spore-bearing plants (the Cryptogams) and an upper kingdom of seed-bearing plants (the Phanerogams). Again, just as the first half of the earth’s story is the age of Invertebrate animals, so it is the age of Cryptogamous plants. So far evolution was always justified in the plant record. But there is a third parallel, of much greater interest. We saw that at one time the evolutionist was puzzled by the clean division of animals into Invertebrate and Vertebrate, and the sudden appearance of the backbone in the chronicle: he was just as much puzzled by the sharp division of our plants into Cryptogams and Phanerogams, and the sudden appearance of the latter on the earth during the Coal-forest period. And the issue has been a fresh and recent triumph for evolution.

Plants are so well preserved in the coal that many years of microscopic study of the remains, and patient putting-together of the crushed and scattered fragments, have shown the Carboniferous plants in quite a new light. Instead of the Coal-forest being a vast assemblage of Cryptogams, upon which the higher type of the Phanerogam is going suddenly to descend from the clouds, it is, to a very great extent, a world of plants that are struggling upward, along many paths, to the higher level. The characters of the Cryptogam and Phanerogam are so mixed up in it that, although the special lines of development are difficult to trace, it is one massive testimony to the evolution of the higher from the lower. The reproductive bodies of the great Lepidodendra are sometimes more like seeds than spores, while both the wood and the leaves of the Sigillaria have features which properly belong to the Phanerogam. In another group (called the Sphenophyllales) the characters of these giant Club-mosses are blended with the characters of the giant Horsetails, and there is ground to think that the three groups have descended from an earlier common ancestor.

Further, it is now believed that a large part of what were believed to be Conifers, suddenly entering from the unknown, are not Conifers at all, but Cordaites. The Cordaites is a very remarkable combination of features that are otherwise scattered among the Cryptogams, Cycads, and Conifers. On the other hand, a very large part of what the geologist had hitherto called “Ferns” have turned out to be seed-bearing plants, half Cycad and half Fern. Numbers of specimens of this interesting group ­the Cycadofilices (cycad-ferns) or Pteridosperms (seed-ferns) ­have been beautifully restored by our botanists. They have afforded a new and very plausible ancestor for the higher trees which come on the scene toward the close of the Coal-forests, while their fern-like characters dispose botanists to think that they and the Ferns may be traced to a common ancestor. This earlier stage is lost in those primitive ages from which not a single leaf has survived in the rocks. We can only say that it is probable that the Mosses, Ferns, Lycopods, etc., arose independently from the primitive level. But the higher and more important development is now much clearer. The Coal-forest is not simply a kingdom of Cryptogams. It is a world of aspiring and mingled types. Let it be subjected to some searching test, some tremendous spell of adversity, and we shall understand the emergence of the higher types out of the luxuriant profusion and confusion of forms.