The camel seems to be traceable to a group of primitive North American Ungulates (Paebrotherium, etc.) in the later Eocene period. The Paebrotherium, a small animal about two feet long, is followed by Pliauchenia, which points toward the llamas and vicunas, and Procamelus, which clearly foreshadows the true camel. In the Pliocene the one branch went southward, to develop into the llamas and vicunas, and the other branch crossed to Asia, to develop into the camels. Since that time they have had no descendants in North America. The primitive giraffe appears suddenly in the later Tertiary deposits of Europe and Asia. The evidence points to an invasion from Africa, and, as the region of development is unknown and unexplored, the evolution of the giraffe remains a matter of speculation. Chevrotains flourished in Europe and North America in the Oligocene, and are still very primitive in structure, combining features of the hog and the ruminants. Primitive deer and oxen begin in the Miocene, and seem to have an earlier representative in certain American animals (Protoceras), of which the male has a pair of blunt outgrowths between the ears. The first true deer are hornless (like the primitive muskdeer of Asia to-day), but by the middle of the Miocene the males have small two-pronged antlers, and as the period proceeds three or four more prongs are added. It is some confirmation of the evolutionary embryonic law that we find the antlers developing in this way in the individual stag to-day. A very curious race of ruminants in the later Tertiary was a large antelope (Sivatherium) with four horns. It had not only the dimensions, but apparently some of the characters, of an elephant. The elephant itself, the last type of the Ungulates, has a clearer line of developments. A chance discovery of fossils in the Fayum district in Egypt led Dr. C. W. Andrews to make a special exploration, and on the remains which he found he has constructed a remarkable story of the evolution of the elephant. [*] It is clear that the elephant was developed in Africa, and a sufficiently complete series of remains has been found to give a good idea of the origin of its most distinctive features. In the Eocene period there lived in the Egyptian region an animal, something like the tapir in size and appearance, which had its second incisors developed into small tusks and--to judge from the nasal opening in the skull--a somewhat prolonged snout. This animal (Moeritherium) only differed from the ordinary primitive Ungulate in these incipient elephantine features. In the later Eocene a larger and more advanced animal, the Palaeomastodon, makes its appearance. Its tusks are larger (five or six inches long), its molars more elephantine, the air-cells at the back of the head more developed. It would look like a small elephant, except that it had a long snout, instead of a flexible trunk,
and a projecting lower jaw on which the snout rested.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. [*] * Of recent years another compromise has been proposed between the Lamarckians and Weismannists. It would say that the efforts of the parent and their effect on the position of the eye--in our case--are not inherited, but might be of use in sheltering an embryonic variation in the direction of a displaced eye. 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.The Archaean continent that we described was being reduced constantly by the wash of rain, the scouring of rivers, and the fretting of the waves on the coast. It is generally thought that these wearing agencies were more violent in early times, but that is disputed, and we will not build on it. In any case, in the course of time millions of tons of matter were scraped off the Archaean continent and laid on the floor of the sea by its rivers. This meant a very serious alteration of pressure or weight on the surface of the globe, and was bound to entail a reaction or restoration of the balance. The rise of the land and formation of mountains used to be ascribed mainly to the cooling and shrinking of the globe of the earth. The skin (crust), it was thought, would become too large for the globe as it shrank, and would wrinkle outwards, or pucker up into mountain-chains. The position of our greater mountain-chains sprawling across half the earth (the Pyrenees to the Himalaya, and the Rocky Mountains to the Andes), seems to confirm this, but the question of the interior of the earth is obscure and disputed, and geologists generally conceive the rise of land and formation of mountains in a different way. They are due probably to the alteration of pressure on the crust in combination with the instability of the interior. The floors of the seas would sink still lower under their colossal burdens, and this would cause some draining of the land-surface. At the same time the heavy pressure below the seas and the lessening of pressure over the land would provoke a reaction. Enormous masses of rock would be forced toward and underneath the land-surface, bending, crumpling, and upheaving it as if its crust were but a leather coat. As a result, masses of land would slowly rise above the plain, to be shaped into hills and valleys by the hand of later time, and fresh surfaces would be dragged out of the deep, enlarging the fringes of the primitive continents, to be warped and crumpled in their turn at the next era of pressure. In point of geological fact, the story of the earth has been one prolonged series of changes in the level of land and water, and in their respective limits. These changes have usually been very gradual, but they have always entailed changes (in climate, etc. ) of the greatest significance in the evolution of life. What was the swampy soil of England in the Carboniferous period is now sometimes thousands of feet beneath us; and what was the floor of a deep ocean over much of Europe and Asia at another time is now to be found on the slopes of lofty Alps, or 20,000 feet above the sea-level in Thibet. Our story of terrestrial life will be, to a great extent, the story of how animals and plants changed their structure in the long series of changes which this endless battle of land and sea brought over the face of the earth. As we have no recognisable remains of the animals and plants of the earliest age, we will not linger over the Archaean rocks. Starting from deep and obscure masses of volcanic matter, the geologist, as he travels up the series of Archaean rocks, can trace only a dim and most unsatisfactory picture of those remote times. Between outpours of volcanic floods he finds, after a time, traces that an ocean and rivers are wearing away the land. He finds seams of carbon among the rocks of the second division of the Archaean (the Keewatin), and deduces from this that a dense sea-weed population already covered the floor of the ocean. In the next division (the Huronian) he finds the traces of extensive ice-action strangely lying between masses of volcanic rock, and sees that thousands of square miles of eastern North America were then covered with an ice-sheet. Then fresh floods of molten matter are poured out from the depths below; then the sea floods the land for a time; and at last it makes its final emergence as the first definitive part of the North American continent, to enlarge, by successive fringes, to the continent of to-day. [*] * I am quoting Professor Coleman's summary of Archaean research in North America (Address to the Geological Section of the British Association, 1909). Europe, as a continent, has had more "ups and downs" than America in the course of geological time. This meagre picture of the battle of land and sea, with interludes of great volcanic activity and even of an ice age, represents nearly all we know of the first half of the world's story from geology. It is especially disappointing in regard to the living population. The very few fossils we find in the upper Archaean rocks are so similar to those we shall discuss in the next chapter that we may disregard them, and the seams of carbon-shales, iron-ore, and limestone, suggest only, at the most, that life was already abundant. We must turn elsewhere for some information on the origin and early development of life. The question of the origin of life I will dismiss with a brief account of the various speculations of recent students of science. Broadly speaking, their views fall into three classes. Some think that the germs of life may have come to the earth from some other body in the universe; some think that life was evolved out of non-living matter in the early ages of the earth, under exceptional conditions which we do not at present know, or can only dimly conjecture; and some think that life is being evolved from non-life in nature to-day, and always has been so evolving. The majority of scientific men merely assume that the earliest living things were no exception to the general process of evolution, but think that we have too little positive knowledge to speculate profitably on the manner of their origin. The first view, that the germs of life may have come to this planet on a meteoric visitor from some other world, as a storm-driven bird may take its parasites to some distant island, is not without adherents to-day. It was put forward long ago by Lord Kelvin and others; it has been revived by the distinguished Swede, Professor Svante Arrhenius. The scientific objection to it is that the more intense (ultra-violet) rays of the sun would frill such germs as they pass through space. But a broader objection, and one that may dispense us from dwelling on it, is that we gain nothing by throwing our problems upon another planet. We have no ground for supposing that the earth is less capable of evolving life than other planets. The second view is that, when the earth had passed through its white-hot stage, great masses of very complex chemicals, produced by the great heat, were found on its surface. There is one complex chemical substance in particular, called cyanogen, which is either an important constituent of living matter, or closely akin to it. Now we need intense heat to produce this substance in the laboratory. May we not suppose that masses of it were produced during the incandescence of the earth, and that, when the waters descended, they passed through a series of changes which culminated in living plasm? Such is the "cyanogen hypothesis" of the origin of life, advocated by able physiologists such as Pfluger, Verworn, and others. It has the merit of suggesting a reason why life may not be evolving from non-life in nature to-day, although it may have so evolved in the Archaean period. Other students suggest other combinations of carbon-compounds and water in the early days. Some suggest that electric action was probably far more intense in those ages; others think that quantities of radium may have been left at the surface. But the most important of these speculations on the origin of life in early times, and one that has the merit of not assuming any essentially different conditions then than we find now, is contained in a recent pronouncement of one of the greatest organic chemists in Europe, Professor Armstrong. He says that such great progress has been made in his science--the science of the chemical processes in living things--that "their cryptic character seems to have disappeared almost suddenly." On the strength of this new knowledge of living matter, he ventures to say that "a series of lucky accidents" could account for the first formation of living things out of non-living matter in Archaean times. Indeed, he goes further. He names certain inorganic substances, and says that the blowing of these into pools by the wind on the primitive planet would set afoot chemical combinations which would issue in the production of living matter. [*] * See his address in Nature, vol. 76, p. 651. For other speculations see Verworn's "General Physiology," Butler Burke's "Origin of Life" (1906), and Dr. Bastian's "Origin of Life" (1911). It is evident that the popular notion that scientific men have declared that life cannot be evolved from non-life is very far astray. This blunder is usually due to a misunderstanding of the dogmatic statement which one often reads in scientific works that "every living thing comes from a living thing." This principle has no reference to remote ages, when the conditions may have been different. It means that to-day, within our experience, the living thing is always born of a living parent. However, even this is questioned by some scientific men of eminence, and we come to the third view. Professor Nageli, a distinguished botanist, and Professor Haeckel, maintain that our experience, as well as the range of our microscopes, is too limited to justify the current axiom. They believe that life may be evolving constantly from inorganic matter. Professor J. A. Thomson also warns us that our experience is very limited, and, for all we know, protoplasm may be forming naturally in our own time. Mr. Butler Burke has, under the action of radium, caused the birth of certain minute specks which strangely imitate the behaviour of bacteria. Dr. Bastian has maintained for years that he has produced living things from non-living matter. In his latest experiments, described in the book quoted, purely inorganic matter is used, and it is previously subjected, in hermetically sealed tubes, to a heat greater than what has been found necessary to kill any germs whatever. Evidently the problem of the origin of life is not hopeless, but our knowledge of the nature of living matter is still so imperfect that we may leave detailed speculation on its origin to a future generation. Organic chemistry is making such strides that the day may not be far distant when living matter will be made by the chemist, and the secret of its origin revealed. For the present we must be content to choose the more plausible of the best-informed speculations on the subject. But while the origin of life is obscure, the early stages of its evolution come fairly within the range of our knowledge. To the inexpert it must seem strange that, whereas we must rely on pure speculation in attempting to trace the origin of life, we can speak with more confidence of those early developments of plants and animals which are equally buried in the mists of the Archaean period. Have we not said that nothing remains of the procession of organisms during half the earth's story but a shapeless seam of carbon or limestone? A simple illustration will serve to justify the procedure we are about to adopt. Suppose that the whole of our literary and pictorial references to earlier stages in the development of the bicycle, the locomotive, or the loom, were destroyed. We should still be able to retrace the phases of their evolution, because we should discover specimens belonging to those early phases lingering in our museums, in backward regions, and elsewhere. They might yet be useful in certain environments into which the higher machines have not penetrated. In the same way, if all the remains of prehistoric man and early civilisation were lost, we could still fairly retrace the steps of the human race, by gathering the lower tribes and races, and arranging them in the order of their advancement. They are so many surviving illustrations of the stages through which mankind as a whole has passed. Just in the same way we may marshal the countless species of animals and plants to-day in such order that they will, in a general way, exhibit to us the age-long procession of life. From the very start of living evolution certain forms dropped out of the onward march, and have remained, to our great instruction, what their ancestors were millions of years ago. People create a difficulty for themselves by imagining that, if evolution is true, all animals must evolve. A glance at our own fellows will show the error of this. Of one family of human beings, as a French writer has said, one only becomes a Napoleon; the others remain Lucien, Jerome, or Joseph. Of one family of animals or trees, some advance in one or other direction; some remain at the original level. There is no "law of progress." The accidents of the world and hereditary endowment impel some onward, and do not impel others. Hence at nearly every great stage in the upward procession through the ages some regiment of plants or animals has dropped out, and it represents to-day the stage of life at which it ceased to progress. In other words, when we survey the line of the hundreds of thousands of species which we find in nature to-day, we can trace, amid their countless variations and branches, the line of organic evolution in the past; just as we could, from actual instances, study the evolution of a British house, from the prehistoric remains in Devonshire to a mansion in Park Lane or a provincial castle. Another method of retracing the lost early chapters in the development of life is furnished by embryology. The value of this method is not recognised by all embryologists, but there are now few authorities who question the substantial correctness of it, and we shall, as we proceed, see some remarkable applications of it. In brief, it is generally admitted that an animal or plant is apt to reproduce, during its embryonic development, some of the stages of its ancestry in past time. This does not mean that a higher animal, whose ancestors were at one time worms, at another time fishes, and at a later time reptiles, will successively take the form of a little worm, a little fish, and a little reptile. The embryonic life itself has been subject to evolution, and this reproduction of ancestral forms has been proportionately disturbed. Still, we shall find that animals will tend, in their embryonic development, to reproduce various structural features which can only be understood as reminiscences of ancestral organs. In the lower animals the reproduction is much less disturbed than in the higher, but even in the case of man this law is most strikingly verified. We shall find it useful sometimes at least in confirming our conclusions as to the ancestry of a particular group. We have, therefore, two important clues to the missing chapters in the story of evolution. Just as the scheme of the evolution of worlds is written broadly across the face of the heavens to-day, so the scheme of the evolution of life is written on the face of living nature; and it is written again, in blurred and broken characters, in the embryonic development of each individual. With these aids we set out to restore the lost beginning of the epic of organic evolution.