From the description of the Tertiary world which we have seen in the last chapter we understand the rapid evolution of the herbivorous Condylarthra. The rich vegetation which spreads over the northern continents, to which they have penetrated, gives them an enormous vitality and fecundity, and they break into groups, as they increase in number, adapted to the different conditions of forest, marsh, or grass-covered plain. Some of them, swelling lazily on the abundant food, and secure for a time in their strength, become the Deinosaurs of their age, mere feeding and breeding machines. They are massive, sluggish, small-brained animals, their strong stumpy limbs terminating in broad five-toed feet. Coryphodon, sometimes as large as an ox, is a typical representative. It is a type fitted only for prosperous days, and these Amblypoda, as they are called, will disappear as soon as the great carnivores are developed.

Another doomed race, or abortive experiment of early mammal life, were the remarkable Deinocerata ("terrible-horned" mammals). They sometimes measured thirteen feet in length, but had little use for brain in the conditions in which they were developed. The brain of the Deinoceras was only one-eighth the size of the brain of a rhinoceros of the same bulk; and the rhinoceros is a poor-brained representative of the modern mammals. To meet the growing perils of their race they seem to have developed three pairs of horns on their long, flat skulls, as we find on them three pairs of protuberances. A late specimen of the group, Tinoceras, had a head four feet in length, armed with these six horns, and its canine teeth were developed into tusks sometimes seven or eight inches in length. They suggest a race of powerful but clumsy and grotesque monsters, making a last stand, and developing such means of protection as their inelastic nature permitted. But the horns seem to have proved a futile protection against the advancing carnivores, and the race was extinguished. The horns may, of course, have been mainly developed by, or for, the mutual butting of the males.

The extinction of these races will remind many readers of a theory on which it is advisable to say a word. It will be remembered that the last of the Deinosaurs and the Ammonites also exhibited some remarkable developments in their last days. These facts have suggested to some writers the idea that expiring races pass through a death-agony, and seem to die a natural death of old age like individuals. The Trilobites are quoted as another instance; and some ingenious writers add the supposed eccentricities of the Roman Empire in its senile decay and a number of other equally unsubstantial illustrations.

There is not the least ground for this fantastic speculation. The destruction of these "doomed races" is as clearly traceable to external causes as is the destruction of the Roman Empire; nor, in fact, did the Roman Empire develop any such eccentricities as are imagined in this superficial theory. What seem to our eye the "eccentricities" and "convulsions" of the Ceratopsia and Deinocerata are much more likely to be defensive developments against a growing peril, but they were as futile against the new carnivores as were the assegais of the Zulus against the European. On the other hand, the eccentricities of many of the later Trilobites—the LATEST Trilobites, it may be noted, were chaste and sober specimens of their race, like the last Roman patricians—and of the Ammonites may very well have been caused by physical and chemical changes in the sea-water. We know from experiment that such changes have a disturbing influence, especially on the development of eggs and larvae; and we know from the geological record that such changes occurred in the periods when the Trilobites and Ammonites perished. In fine, the vast majority of extinct races passed through no "convulsions" whatever. We may conclude that races do not die; they are killed.

The extinction of these races of the early Condylarthra, and the survival of those races whose descendants share the earth with us to-day, are quite intelligible. The hand of natural selection lay heavy on the Tertiary herbivores. Apart from overpopulation, forcing groups to adapt themselves to different regions and diets, and apart from the geological disturbances and climatic changes which occurred in nearly every period, the shadow of the advancing carnivores was upon them. Primitive but formidable tigers, wolves, and hyenas were multiplying, and a great selective struggle set in. Some groups shrank from the battle by burrowing underground like the rabbit; some, like the squirrel or the ape, took refuge in the trees; some, like the whale and seal, returned to the water; some shrank into armour, like the armadillo, or behind fences of spines, like the hedgehog; some, like the bat, escaped into the air. Social life also was probably developed at this time, and the great herds had their sentinels and leaders. But the most useful qualities of the large vegetarians, which lived on grass and leaf, were acuteness of perception to see the danger, and speed of limb to escape it. In other words, increase of brain and sense-power and increase of speed were the primary requisites. The clumsy early Condylarthra failed to meet the tests, and perished; the other branches of the race were more plastic, and, under the pressure of a formidable enemy, were gradually moulded into the horse, the deer, the ox, the antelope, and the elephant.

We can follow the evolution of our mammals of this branch most easily by studying the modification of the feet and limbs. In a running attitude—the experiment may be tried—the weight of the body is shifted from the flat sole of the foot, and thrown upon the toes, especially the central toes. This indicates the line of development of the Ungulates (hoofed animals) in the struggle of the Tertiary Era. In the early Eocene we find the Condylarthra (such as Phenacodus) with flat five-toed feet, and such a mixed combination of characters that they "might serve very well for the ancestors of all the later Ungulata" (Woodward). We then presently find this generalised Ungulate branching into three types, one of which seems to be a patriarchal tapir, the second is regarded as a very remote ancestor of the horse, and the third foreshadows the rhinoceros. The feet have now only three or four toes; one or two of the side-toes have disappeared. This evolution, however, follows two distinct lines. In one group of these primitive Ungulates the main axis of the limb, or the stress of the weight, passes through the middle toe. This group becomes the Perissodactyla ("odd-toed" Ungulates) of the zoologist, throwing out side-branches in the tapir and the rhinoceros, and culminating in the one-toed horse. In the other line, the Artiodactyla (the "even-toed" or cloven-hoofed Ungulates), the main axis or stress passes between the third and fourth toes, and the group branches into our deer, oxen, sheep, pigs, camels, giraffes, and hippopotamuses. The elephant has developed along a separate and very distinctive line, as we shall see, and the hyrax is a primitive survivor of the ancestral group.

Thus the evolutionist is able to trace a very natural order in the immense variety of our Ungulates. He can follow them in theory as they slowly evolve from their primitive Eocene ancestor according to their various habits and environments; he has a very rich collection of fossil remains illustrating the stages of their development; and in the hyrax (or "coney") he has one more of those living fossils, or primitive survivors, which still fairly preserve the ancestral form. The hyrax has four toes on the front foot and three on the hind foot, and the feet are flat. Its front teeth resemble those of a rodent, and its molars those of the rhinoceros. In many respects it is a most primitive and generalised little animal, preserving the ancestral form more or less faithfully since Tertiary days in the shelter of the African Continent.

The rest of the Ungulates continued to develop through the Tertiary, and fortunately we are enabled to follow the development of two of the most interesting of them, the horse and the elephant, in considerable detail. As I said above, the primitive Ungulate soon branches into three types which dimly foreshadow the tapir, the horse, and the rhinoceros, the three forms of the Perissodactyl. The second of these types is the Hyracotherium. It has no distinct equine features, and is known only from the skull, but the authorities regard it as the progenitor (or representative of the progenitors) of the horse-types. In size it must have been something like the rabbit or the hyrax. Still early in the Eocene, however, we find the remains of a small animal (Eohippus), about the size of a fox, which is described as "undoubtedly horse-like." It had only three toes on its hind feet, and four on its front feet; though it had also a splint-bone, representing the shrunken and discarded fifth toe, on its fore feet. Another form of the same period (Protorohippus) shows the central of the three toes on the hind foot much enlarged, and the lateral toes shrinking. The teeth, and the bones and joints of the limbs, are also developing in the direction of the horse.

In the succeeding geological period, the Oligocene, we find several horse-types in which the adaptation of the limbs to running on the firm grassy plains and of the teeth to eating the grass continues. Mesohippus has lost the fourth toe of the fore foot, which is now reduced to a splintbone, and the lateral toes of its hind foot are shrinking. In the Miocene period there is a great development of the horse-like mammals. We have the remains of more than forty species, some continuing the main line of development on the firm and growing prairies of the Miocene, some branching into the softer meadows or the forests, and giving rise to types which will not outlive the Tertiary. They have three toes on each foot, and have generally lost even the rudimentary trace of the fourth toe. In most of them, moreover, the lateral toes—except in the marsh-dwelling species, with spreading feet—scarcely touch the ground, while the central toe is developing a strong hoof. The leg-bones are longer, and have a new type of joint; the muscles are concentrated near the body. The front teeth are now chopping incisors, and the grinding teeth approach those of the modern horse in the distribution of the enamel, dentine, and cement. They are now about the size of a donkey, and must have had a distinctly horsy appearance, with their long necks and heads and tapering limbs. One of them, Merychippus, was probably in the direct line of the evolution of the horse. From Hipparion some of the authorities believe that the zebras may have been developed. Miohippus, Protohippus, and Hypohippus, varying in size from that of a sheep to that of a donkey, are other branches of this spreading family.

In the Pliocene period the evolution of the main stem culminates in the appearance of the horse, and the collateral branches are destroyed. Pliohippus is a further intermediate form. It has only one toe on each foot, with two large splint bones, but its hoof is less round than that of the horse, and it differs in the shape of the skull and the length of the teeth. The true horse (Equus) at length appears, in Europe and America, before the close of the Tertiary period. As is well known, it still has the rudimentary traces of its second and fourth toes in the shape of splint bones, and these bones are not only more definitely toe-shaped in the foal before birth, but are occasionally developed and give us a three-toed horse.

From these successive remains we can confidently picture the evolution, during two or three million years, of one of our most familiar mammals. It must not, of course, be supposed that these fossil remains all represent "ancestors of the horse." In some cases they may very well do so; in others, as we saw, they represent sidebranches of the family which have become extinct. But even such successive forms as the Eohippus, Mesohippus, Miohippus, and Pliohippus must not be arranged in a direct line as the pedigree of the horse. The family became most extensive in the Miocene, and we must regard the casual fossil specimens we have discovered as illustrations of the various phases in the development of the horse from the primitive Ungulate. When we recollect what we saw in an earlier chapter about the evolution of grassy plains and the successive rises of the land during the Tertiary period, and when we reflect on the simultaneous advance of the carnivores, we can without difficulty realise this evolution of our familiar companion from a hyrax-like little animal of two million years ago.

We have not in many cases so rich a collection of intermediate forms as in the case of the horse, but our fossil mammals are numerous enough to suggest a similar development of all the mammals of to-day. The primitive family which gave birth to the horse also gave us, as we saw, the tapir and the rhinoceros. We find ancestral tapirs in Europe and America during the Tertiary period, but the later cold has driven them to the warm swamps of Brazil and Malaysia. The rhinoceros has had a long and interesting history. From the primitive Hyrochinus of the Eocene, in which it is dimly foreshadowed, we pass to a large and varied family in the later periods. In the Oligocene it spreads into three great branches, adapted, respectively, to life on the elevated lands, the lowlands, and the water. The upland type (Hyracodon) was a light-limbed running animal, well illustrating the close relation to the horse. The aquatic representative (Metamynodon) was a stumpy and bulky animal. The intermediate lowland type was probably the ancestor of the modern animal. All three forms were yet hornless. In the Miocene the lowland type (Leptaceratherium, Aceratherium, etc.) develops vigorously, while the other branches die. The European types now have two horns, and in one of the American species (Diceratherium) we see a commencement of the horny growths from the skull. We shall see later that the rhinoceros continued in Europe even during the severe conditions of the glacial period, in a branch that developed a woolly coat.

There were also in the early Tertiary several sidebranches of the horse-tapir-rhinoceros family. The Palaeotheres were more or less between the horse and the tapir in structure; the Anoplotheres between the tapir and the ruminant. A third doomed branch, the Titanotheres, flourished vigorously for a time, and begot some strange and monstrous forms (Brontops, Titanops, etc.). In the larger specimens the body was about fourteen feet long, and stood ten feet from the ground. The long, low skull had a pair of horns over the snout. They perished like the equally powerful but equally sluggish and stupid Deinocerata. The Tertiary was an age of brain rather than of brawn. As compared with their early Tertiary representatives' some of our modern mammals have increased seven or eight-fold in brain-capacity.

While the horses and tapirs and rhinoceroses were being gradually evolved from the primitive types, the Artiodactyl branch of the Ungulates—the pigs, deer, oxen, etc.—were also developing. We must dismiss them briefly. We saw that the primitive herbivores divided early in the Eocene into the "odd-toed" and "even-toed" varieties; the name refers, it will be remembered, not to the number of toes, but to the axis of stress. The Artiodactyl group must have quickly branched in turn, as we find very primitive hogs and camels before the end of the Eocene. The first hog-like creature (Homacodon) was much smaller than the hog of to-day, and had strong canine teeth, but in the Oligocene the family gave rise to a large and numerous race, the Elotheres. These "giant-pigs," as they have been called, with two toes on each foot, flourished vigorously for a time in Europe and America, but were extinguished in the Miocene, when the true pigs made their appearance. Another doomed race of the time is represented by the Hyopotamus, an animal between the pig and the hippopotamus; and the Oreodontids, between the hog and the deer, were another unsuccessful branch of the early race. The hippopotamus itself was widespread in Europe, and a familiar form in the rivers of Britain, in the latter part of the Tertiary.

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.

*See this short account, "Guide to the Elephants in the British Museum," 1908.

Up to the beginning of the Miocene, Africa was, as we saw, cut off from Europe and Asia by the sea which stretched from Spain to India. Then the land rose, and the elephant passed by the new tracts into the north. Its next representative, Tetrabelodon, is found in Asia and Europe, as well as North Africa. The frame is as large as that of a medium-sized elephant, and the increase of the air-cells at the back of the skull shows that an increased weight has to be sustained by the muscles of the neck. The nostrils are shifted further back. The tusks are from twenty to thirty inches long, and round, and only differ from those of the elephant in curving slightly downward, The chin projects as far as the tusks. The neck is shorter and thicker, and, as the animal increases in height, we can understand that the long snout—possibly prehensile at its lower end—is necessary for the animal to reach the ground. But the snout still lies on the projecting lower jaw, and is not a trunk. Passing over the many collateral branches, which diverge in various directions, we next kind that the chin is shortening (in Tetrabelodon longirostris), and, through a long series of discovered intermediate forms, we trace the evolution of the elephant from the mastodon. The long supporting skin disappears, and the enormous snout becomes a flexible trunk. Southern Asia seems to have been the province of this final transformation, and we have remains of some of these primitive elephants with tusks nine and a half feet long. A later species, which wandered over Central and Southern Europe before the close of the Tertiary, stood fifteen feet high at the shoulder, while the mammoth, which superseded it in the days of early man, had at times tusks more than ten feet in length.

It is interesting to reflect that this light on the evolution of one of our most specialised mammals is due to the chance opening of the soil in an obscure African region. It suggests to us that as geological exploration is extended, many similar discoveries may be made. The slenderness of the geological record is a defect that the future may considerably modify.

From this summary review of the evolution of the Ungulates we must now pass to an even briefer account of the evolution of the Carnivores. The evidence is less abundant, but the characters of the Carnivores consist so obviously of adaptations to their habits and diet that we have little difficulty in imagining their evolution. Their early Eocene ancestors, the Creodonts, gave rise in the Eocene to forms which we may regard as the forerunners of the cat-family and dog-family, to which most of our familiar Carnivores belong. Patriofelis, the "patriarchal cat," about five or six feet in length (without the tail), curiously combines the features of the cat and the seal-family. Cyonodon has a wolf-like appearance, and Amphicyon rather suggests the fox. Primitive weasels, civets, and hyaenas appear also in the Eocene. The various branches of the Carnivore family are already roughly represented, but it is an age of close relationships and generalised characters.

In the Miocene we find the various groups diverging still further from each other and from the extinct stocks. Definite wolves and foxes abound in America, and the bear, civet, and hyaena are represented in Europe, together with vague otter-like forms. The dog-family seems to have developed chiefly in North America. As in the case of the Ungulates, we find many strange side-branches which flourished for a time, but are unknown to-day. Machoerodus, usually known as "the sabre-toothed tiger," though not a tiger, was one of the most formidable of these transitory races. Its upper canine teeth (the "sabres") were several inches in length, and it had enormously distensible jaws to make them effective. The great development of such animals, with large numbers of hyaenas, civets, wolves, bears, and other Carnivores, in the middle and later Tertiary was probably the most effective agency in the evolution of the horse and deer and the extinction of the more sluggish races. The aquatic branch of the Carnivores (seals, walruses, etc.) is little represented in the Tertiary record. We saw, however, that the most primitive representatives of the elephant-stock had also some characters of the seal, and it is thought that the two had a common origin.

The Moeritherium was a marsh-animal, and may very well have been cousin to the branch of the family which pushed on to the seas, and developed its fore limbs into paddles.

The Rodents are represented in primitive form early in the Eocene period. The teeth are just beginning to show the characteristic modification for gnawing. A large branch of the family, the Tillodonts, attained some importance a little later. They are described as combining the head and claws of a bear with the teeth of a rodent and the general characters of an ungulate. In the Oligocene we find primitive squirrels, beavers, rabbits, and mice. The Insectivores also developed some of the present types at an early date, and have since proved so unprogressive that some regard them as the stock from which all the placental mammals have arisen.

The Cetacea (whales, porpoises, etc.) are already represented in the Eocene by a primitive whale-like animal (Zeuglodon) of unknown origin. Some specimens of it are seventy feet in length. It has large teeth, sometimes six inches long, and is clearly a terrestrial mammal that has returned to the waters. Some forms even of the modern whale develop rudimentary teeth, and in all forms the bony structure of the fore limbs and degenerate relic of a pelvis and back limbs plainly tell of the terrestrial origin. Dolphins appear in the Miocene.

Finally, the Edentates (sloths, anteaters, and armadilloes) are represented in a very primitive form in the early Eocene. They are then barely distinguishable from the Condylarthra and Creodonta, and seem only recently to have issued from a common ancestor with those groups. In the course of the Tertiary we find them—especially in South America, which was cut off from the North and its invading Carnivores during the Eocene and Miocene—developed into large sloths, armadilloes, and anteaters. The reconnection with North America in the Pliocene allowed the northern animals to descend, but gigantic sloths (Megatherium) and armadilloes (Glyptodon) flourished long afterwards in South America. The Megatherium attained a length of eighteen feet in one specimen discovered, and the Glyptodon often had a dorsal shield (like that of the armadillo) from six to eight feet long, and, in addition, a stoutly armoured tail several feet long.

The richness and rapidity of the mammalian development in the Tertiary, of which this condensed survey will convey some impression, make it impossible to do more here than glance over the vast field and indicate the better-known connections. It will be seen that evolution not only introduces a lucid order and arrangement into our thousands of species of living and fossil mammals, but throws an admirable light on the higher animal world of our time. The various orders into which the zoologist puts our mammals are seen to be the branches of a living tree, approaching more and more closely to each other in early Tertiary times, in spite of the imperfectness of the geological record. We at last trace these diverging lines to a few very primitive, generalised, patriarchal groups, which in turn approach each other very closely in structure, and plainly suggest a common Cretaceous ancestor. Whether that common ancestor was an Edentate, an Insectivore, or Creodont, or something more primitive than them all, is disputed. But the divergence of nearly all the lines of our mammal world from those patriarchal types is admirably clear. In the mutual struggle of carnivore and herbivore, in adaptation to a hundred different environments (the water, the land, and the air, the tree, the open plain, the underground, the marsh, etc.) and forms of diet, we find the descendants of these patriarchal animals gradually developing their distinctive characters. Then we find the destructive agencies of living and inorganic nature blotting out type after type, and the living things that spread over the land in the later Tertiary are found to be broadly identical with the living things of to-day. The last great selection, the northern Ice-Age, will give the last touches of modernisation.



CHAPTER XVIII. THE EVOLUTION OF MAN

We have reserved for a closer inquiry that order of the placental mammals to which we ourselves belong, and on which zoologists have bestowed the very proper and distinguishing name of the Primates. Since the days of Darwin there has been some tendency to resent the term "lower animals," which man applies to his poorer relations. But, though there is no such thing as an absolute standard by which we may judge the "higher" or "lower" status of animals or plants, the extraordinary power which man has by his brain development attained over both animate and inanimate nature fully justifies the phrase. The Primate order is, therefore, of supreme interest as the family that gave birth to man, and it is important to discover the agencies which impelled some primitive member of it to enter upon the path which led to this summit of organic nature.

The order includes the femurs, a large and primitive family with ape-like features—the Germans call them "half-apes"—the monkeys, the man-like apes, and man. This classification according to structure corresponds with the successive appearance of the various families in the geological record. The femurs appear in the Eocene; the monkeys, and afterwards the apes, in the Miocene, the first semi-human forms in the Pleistocene, though they must have been developed before this. It is hardly necessary to say that science does not regard man as a descendant of the known anthropoid apes, or these as descended from the monkeys. They are successive types or phases of development, diverging early from each other. Just as the succeeding horse-types of the record are not necessarily related to each other in a direct line, yet illustrate the evolution of a type which culminates in the horse, so the spreading and branching members of the Primate group illustrate the evolution of a type of organism which culminates in man. The particular relationship of the various families, living and dead, will need careful study.

That there is a general blood-relationship, and that man is much more closely related to the anthropoid apes than to any of the lower Primates, is no longer a matter of controversy. In Rudolph Virchow there died, a few years ago, the last authoritative man of science to express any doubt about it. There are, however, non-scientific writers who, by repeating the ambiguous phrase that it is "only a theory," convey the impression to inexpert readers that it is still more or less an open question. We will therefore indicate a few of the lines of evidence which have overcome the last hesitations of scientific men, and closed the discussion as to the fact.

The very close analogy of structure between man and the ape at once suggests that they had a common ancestor. There are cases in which two widely removed animals may develop a similar organ independently, but there is assuredly no possibility of their being alike in all organs, unless by common inheritance. Yet the essential identity of structure in man and the ape is only confirmed by every advance of science, and would of itself prove the common parentage. Such minor differences as there are between man and the higher ape—in the development of the cerebrum, the number of the teeth or ribs, the distribution of the hair, and so on—are quite explicable when we reflect that the two groups must have diverged from each other more than a million years ago.

Examining the structure of man more closely, we find this strong suggestion of relationship greatly confirmed. It is now well known that the human body contains a number of vestigial "organs"—organs of no actual use, and only intelligible as vestiges of organs that were once useful. Whatever view we take of the origin of man, each organ in his frame must have a meaning; and, as these organs are vestigial and useless even in the lowest tribes of men, who represent primitive man, they must be vestiges of organs that were of use in a remote pre-human ancestor. The one fact that the ape has the same vestigial organs as man would, on a scientific standard of evidence, prove the common descent of the two. But these interesting organs themselves point back far earlier than a mixed ape-human ancestor in many cases.

The shell of cartilage which covers the entrance to the ear—the gristly appendage which is popularly called the ear—is one of the clearest and most easily recognised of these organs. The "ear" of a horse or a cat is an upright mobile shell for catching the waves of sound. The human ear has the appearance of being the shrunken relic of such an organ, and, when we remove the skin, and find seven generally useless muscles attached to it, obviously intended to pull the shell in all directions (as in the horse), there can be no doubt that the external ear is a discarded organ, a useless legacy from an earlier ancestor. In cases where it has been cut off it was found that the sense of hearing was scarcely, if at all, affected. Now we know that it is similarly useless in all tribes of men, and must therefore come from a pre-human ancestor. It is also vestigial in the higher apes, and it is only when we descend to the lower monkeys and femurs that we see it approaching its primitive useful form. One may almost say that it is a reminiscence of the far-off period when, probably in the early Tertiary, the ancestors of the Primates took to the trees. The animals living on the plain needed acute senses to detect the approach of their prey or their enemies; the tree-dweller found less demand on his sense of hearing, the "speaking-trumpet" was discarded, and the development of the internal ear proceeded on the higher line of the perception of musical sounds.

We might take a very large number of parts of the actual human body, and discover that they are similar historical or archaeological monuments surviving in a modern system, but we have space only for a few of the more conspicuous.

The hair on the body is a vestigial organ, of actual use to no race of men, an evident relic of the thick warm coat of an earlier ancestor. It in turn recalls the dwellers in the primeval forest. In most cases—not all, because the wearing of clothes for ages has modified this feature—it will be found that the hairs on the arm tend upward from the wrist to the elbow, and downward from the shoulder to the elbow. This very peculiar feature becomes intelligible when we find that some of the apes also have it, and that it has a certain use in their case. They put their hands over their heads as they sit in the trees during ram, and in that position the sloping hair acts somewhat like the thatched roof of a cottage.

Again, it will be found that in the natural position of standing we are not perfectly flat-footed, but tend to press much more on the outer than on the inner edge of the foot. This tendency, surviving after ages of living on the level ground, is a lingering effect of the far-off arboreal days.

A more curious reminiscence is seen in the fact that the very young infant, flabby and powerless as it is in most of its muscles, is so strong in the muscles of the hand and arm that it can hang on to a stick by its hands, and sustain the whole weight of its body, for several minutes. Finally, our vestigial tail—for we have a tail comparable to that of the higher apes—must be mentioned. In embryonic development the tail is much longer than the legs, and some children are born with a real tail, which they move as the puppy does, according to their emotional condition. Other features of the body point back to an even earlier stage. The vermiform appendage—in which some recent medical writers have vainly endeavoured to find a utility—is the shrunken remainder of a large and normal intestine of a remote ancestor. This interpretation of it would stand even if it were found to have a certain use in the human body. Vestigial organs are sometimes pressed into a secondary use when their original function has been lost. The danger of this appendage in the human body to-day is due to the fact that it is a blind alley leading off the alimentary canal, and has a very narrow opening. In the ape the opening is larger, and, significantly enough, it is still larger in the human foetus. When we examine some of the lower mammals we discover the meaning of it. It is in them an additional storage chamber in the alimentary system. It is believed that a change to a more digestible diet has made this additional chamber superfluous in the Primates, and the system is slowly suppressing it.

Other reminiscences of this earlier phase are found in the many vestigial muscles which are found in the body to-day. The head of the quadruped hangs forward, and is held by powerful muscles and ligaments in the neck. We still have the shrunken remainder of this arrangement. Other vestigial muscles are found in the forehead, the scalp, the nose—many people can twitch the nostrils and the scalp—and under the skin in many parts of the body. These are enfeebled remnants of the muscular coat by which the quadruped twitches its skin, and drives insects away. A less obvious feature is found by the anatomist in certain blood-vessels of the trunk. As the blood flows vertically in a biped and horizontally in a quadruped, the arrangement of the valves in the blood-vessels should be different in the two cases; but it is the same in us as in the quadruped. Another trace of the quadruped ancestor is found in the baby. It walks "on all fours" so long, not merely from weakness of the limbs, but because it has the spine of a quadruped.

A much more interesting fact, but one less easy to interpret, is that the human male has, like the male ape, organs for suckling the young. That there are real milk-glands, usually vestigial, underneath the teats in the breast of the boy or the man is proved by the many known cases in which men have suckled the young. Several friends of the present writer have seen this done in India and Ceylon by male "wet-nurses." As there is no tribe of men or species of ape in which the male suckles the young normally, we seem to be thrown back once more upon an earlier ancestor. The difficulty is that we know of no mammal of which both parents suckle the young, and some authorities think that the breasts have been transferred to the male by a kind of embryonic muddle. That is difficult to believe, as no other feature has ever been similarly transferred to the opposite sex. In any case the male breasts are vestigial organs. Another peculiarity of the mammary system is that sometimes three, four, or five pairs of breasts appear in a woman (and several have been known even in a man). This is, apparently, an occasional reminiscence of an early mammal ancestor which had large litters of young and several pairs of breasts.

But there are features of the human body which recall an ancestor even earlier than the quadruped. The most conspicuous of these is the little fleshy pad at the inner corner of each eye. It is a common feature in mammals, and is always useless. When, however, we look lower down in the animal scale we find that fishes and reptiles (and birds) have a third eyelid, which is drawn across the eye from this corner. There is little room to doubt that the little fleshy vestige in the mammal's eye is the shrunken remainder of the lateral eyelid of a remote fish-ancestor.

A similar reminiscence is found in the pineal body, a small and useless object, about the size and shape of a hazel-nut, in the centre of the brain. When we examine the reptile we find a third eye in the top of the head. The skin has closed over it, but the skull is still, in many cases, perforated as it is for the eyes in front. I have seen it standing out like a ball on the head of a dead crocodile, and in the living tuatara—the very primitive New Zealand lizard—it still has a retina and optic nerve. As the only animal in nature to-day with an eye in this position (the Pyrosome, a little marine animal of the sea-squirt family) is not in the line of reptile and mammal ancestry, it is difficult to locate the third eye definitely. But when we find the skin closing over it in the amphibian and reptile, then the bone, and then see it gradually atrophying and being buried under the growing brain, we must refer it to some early fish-ancestor. This ancestor, we may recall, is also reflected for a time in the gill-slits and arches, with their corresponding fish-like heart and blood-vessels, during man's embryonic development, as we saw in a former chapter.

These are only a few of the more conspicuous instances of vestigial structures in man. Metchnikoff describes about a hundred of them. Even if there were no remains of primitive man pointing in the direction of a common ancestry with the ape, no lower types of men in existence with the same tendency, no apes found in nature to-day with a structure so strikingly similar to that of man, and no fossil records telling of the divergence of forms from primitive groups in past time, we should be forced to postulate the evolution of man in order to explain his actual features. The vestigial structures must be interpreted as we interpret the buttons on the back of a man's coat. They are useless reminiscences of an age in which they were useful. When their witness to the past is supported by so many converging lines of evidence it becomes irresistible. I will add only one further testimony which has been brought into court in recent years.

The blood consists of cells, or minute disk-shaped corpuscles, floating in a watery fluid, or serum. It was found a few years ago, in the course of certain experiments in mixing the blood of animals, that the serum of one animal's blood sometimes destroyed the cells of the other animal's blood, and at other times did not. When the experiments were multiplied, it was found that the amount of destructive action exercised by one specimen of blood upon another depended on the nearness or remoteness of relationship between the animals. If the two are closely related, there is no disturbance when their blood is mixed; when they are not closely related, the serum of one destroys the cells of the other, and the intensity of the action is in proportion to their remoteness from each other. Another and more elaborate form of the experiment was devised, and the law was confirmed. On both tests it was found by experiment that the blood of man and of the anthropoid ape behaved in such a way as to prove that they were closely related. The blood of the monkey showed a less close relationship—a little more remote in the New World than in the Old World monkeys; and the blood of the femur showed a faint and distant relationship.

The FACT of the evolution of man and the apes from a common ancestor is, therefore, outside the range of controversy in science; we are concerned only to retrace the stages of that evolution, and the agencies which controlled it. Here, unfortunately, the geological record gives us little aid. Tree-dwelling animals are amongst the least likely to be buried in deposits which may preserve their bones for ages. The distribution of femur and ape remains shows that the order of the Primates has been widespread and numerous since the middle of the Tertiary Era, yet singularly few remains of the various families have been preserved.

Hence the origin of the Primates is obscure. They are first foreshadowed in certain femur-like forms of the Eocene period, which are said in some cases (Adapis) to combine the characters of pachyderms and femurs, and in others (Anaptomorphus) to unite the features of Insectivores and femurs. Perhaps the more common opinion is that they were evolved from a branch of the Insectivores, but the evidence is too slender to justify an opinion. It was an age when the primitive placental mammals were just beginning to diverge from each other, and had still many features in common. For the present all we can say is that in the earliest spread of the patriarchal mammal race one branch adopted arboreal life, and evolved in the direction of the femurs and the apes. The generally arboreal character of the Primates justifies this conclusion.

In the Miocene period we find a great expansion of the monkeys. These in turn enter the scene quite suddenly, and the authorities are reduced to uncertain and contradictory conjectures as to their origin. Some think that they develop not from the femurs, but along an independent line from the Insectivores, or other ancestors of the Primates. We will not linger over these early monkeys, nor engage upon the hopeless task of tracing their gradual ramification into the numerous families of the present age. It is clear only that they soon divided into two main streams, one of which spread into the monkeys of America and the other into the monkeys of the Old World. There are important anatomical differences between the two. The monkeys remained in Central and Southern Europe until near the end of the Tertiary. Gradually we perceive that the advancing cold is driving them further south, and the monkeys of Gibraltar to-day are the diminished remnant of the great family that had previously wandered as far as Britain and France.

A third wave, also spreading in the Miocene, equally obscure in its connection with the preceding, introduces the man-like apes to the geologist. Primitive gibbons (Pliopithecus and Pliobylobates), primitive chimpanzees (Palaeopithecus), and other early anthropoid apes (Oreopithecus, Dryopithecus, etc.), lived in the trees of Southern Europe in the second part of the Tertiary Era. They are clearly disconnected individuals of a large and flourishing family, but from the half-dozen specimens we have yet discovered no conclusion can be drawn, except that the family is already branching into the types of anthropoid apes which are familiar to us.

Of man himself we have no certain and indisputable trace in the Tertiary Era. Some remains found in Java of an ape-man (Pithecanthropus), which we will study later, are now generally believed, after a special investigation on the spot, to belong to the Pleistocene period. Yet no authority on the subject doubts that the human species was evolved in the Tertiary Era, and very many, if not most, of the authorities believe that we have definite proof of his presence. The early story of mankind is gathered, not so much from the few fragments of human remains we have, but from the stone implements which were shaped by his primitive intelligence and remain, almost imperishable, in the soil over which he wandered. The more primitive man was, the more ambiguous would be the traces of his shaping of these stone implements, and the earliest specimens are bound to be a matter of controversy. It is claimed by many distinguished authorities that flints slightly touched by the hand of man, or at least used as implements by man, are found in abundance in England, France, and Germany, and belong to the Pliocene period. Continental authorities even refer some of them to the Miocene and the last part of the Oligocene.

The question whether an implement-using animal, which nearly all would agree to regard as in some degree human, wandered over what is now the South of England (Kent, Essex, Dorsetshire, etc.) as many hundred thousand years ago as this claim would imply, is certainly one of great interest. But there would be little use in discussing here the question of the "Eoliths," as these disputed implements are called. A very keen controversy is still being conducted in regard to them, and some of the highest authorities in England, France, and Germany deny that they show any trace of human workmanship or usage. Although they have the support of such high authorities as Sir J. Prestwich, Sir E. Ray Lankester, Lord Avebury, Dr. Keane, Dr. Blackmore, Professor Schwartz, etc., they are one of those controverted testimonies on which it would be ill-advised to rely in such a work as this.

We must say, then, that we have no undisputed traces of man in the Tertiary Era. The Tertiary implements which have been at various times claimed in France, Italy, and Portugal are equally disputed; the remains which were some years ago claimed as Tertiary in the United States are generally disallowed; and the recent claims from South America are under discussion. Yet it is the general feeling of anthropologists that man was evolved in the Tertiary Era. On the one hand, the anthropoid apes were highly developed by the Miocene period, and it would be almost incredible that the future human stock should linger hundreds of thousands of years behind them. On the other hand, when we find the first traces of man in the Pleistocene, this development has already proceeded so far that its earlier phase evidently goes back into the Tertiary. Let us pass beyond the Tertiary Era for a moment, and examine the earliest and most primitive remains we have of human or semi-human beings.

The first appearance of man in the chronicle of terrestrial life is a matter of great importance and interest. Even the least scientific of readers stands, so to say, on tiptoe to catch a first glimpse of the earliest known representative of our race, and half a century of discussion of evolution has engendered a very wide interest in the early history of man. [*]

* A personal experience may not be without interest in this connection. Among the many inquiries directed to me in regard to evolution I received, in one month, a letter from a negro in British Guiana and an extremely sensible query from an inmate of an English asylum for the insane! The problem that beset the latter of the two was whether the Lemuranda preceded the Lemurogona in Eocene times. He had found a contradiction in the statements of two scientific writers.

Fortunately, although these patriarchal bones are very scanty—two teeth, a thigh-bone, and the skull-cap—we are now in a position to form some idea of the nature of their living owner. They have been subjected to so searching a scrutiny and discussion since they were found in Java in 1891 and 1892 that there is now a general agreement as to their nature. At first some of the experts thought that they were the remains of an abnormally low man, and others that they belonged to an abnormally high ape. The majority held from the start that they belonged to a member of a race almost midway between the highest family of apes and the lowest known tribe of men, and therefore fully merited the name of "Ape-Man" (Pithecanthropus). This is now the general view of anthropologists.

The Ape-Man of Java was in every respect entitled to that name. The teeth suggest a lower part of the face in which the teeth and lips projected more than in the most ape-like types of Central Africa. The skull-cap has very heavy ridges over the eyes and a low receding forehead, far less human than in any previously known prehistoric skull. The thigh-bone is very much heavier than any known human femur of the same length, and so appreciably curved that the owner was evidently in a condition of transition from the semi-quadrupedal crouch of the ape to the erect attitude of man. The Ape-Man, in other words, was a heavy, squat, powerful, bestial-looking animal; of small stature, but above the pygmy standard; erect in posture, but with clear traces of the proneness of his ancestor; far removed from the highest ape in brainpower, but almost equally far removed from the lowest savage that is known to us. We shall see later that there is some recent criticism, by weighty authorities, of the earlier statements in regard to the brain of primitive man. This does not apply to the Ape-Man of Java. The average cranial capacity (the amount of brain-matter the skull may contain) of the chimpanzees, the highest apes, is about 600 cubic centimetres. The average cranial capacity of the lowest races of men, of moderate stature, is about 1200. And the cranial capacity of Ape-Man was about 900

It is immaterial whether or no these bones belong to the same individual. If they do not, we have remains of two or three individuals of the same intermediate species. Nor does it matter whether or no this early race is a direct ancestor of the later races of men, or an extinct offshoot from the advancing human stock. It is, in either case, an illustration of the intermediate phase between the ape and man The more important tasks are to trace the relationship of this early human stock to the apes, and to discover the causes of its superior evolution.

The first question has a predominantly technical interest, and the authorities are not agreed in replying to it. We saw that, on the blood-test, man showed a very close relationship to the anthropoid apes, a less close affinity to the Old World monkeys, a more remote affinity to the American monkeys, and a very faint and distant affinity to the femurs. A comparison of their structures suggests the same conclusion. It is, therefore, generally believed that the anthropoid apes and man had a common ancestor in the early Miocene or Oligocene, that this group was closely related to the ancestral group of the Old World monkeys, and that all originally sprang from a primitive and generalised femur-group. In other words, a branch of the earliest femur-like forms diverges, before the specific femur-characters are fixed, in the direction of the monkey; in this still vague and patriarchal group a branch diverges, before the monkey-features are fixed, in the direction of the anthropoids; and this group in turn spreads into a number of types, some of which are the extinct apes of the Miocene, four become the gorilla, chimpanzee, orang, and gibbon of to-day, and one is the group that will become man. To put it still more precisely, if we found a whole series of remains of man's ancestors during the Tertiary, we should probably class them, broadly, as femur-remains in the Eocene, monkey-remains in the Oligocene, and ape-remains in the Miocene. In that sense only man "descends from a monkey."

The far more important question is: How did this one particular group of anthropoid animals of the Miocene come to surpass all its cousins, and all the rest of the mammals, in brain-development? Let us first rid the question of its supposed elements of mystery and make of it a simple problem. Some imagine that a sudden and mysterious rise in intelligence lifted the progenitor of man above its fellows. The facts very quickly dispel this illusion. We may at least assume that the ancestor of man was on a level with the anthropoid ape in the Miocene period, and we know from their skulls that the apes were as advanced then as they are now. But from the early Miocene to the Pleistocene is a stretch of about a million years on the very lowest estimate. In other words, man occupied about a million years in travelling from the level of the chimpanzee to a level below that of the crudest savage ever discovered. If we set aside the Java man, as a possible survivor of an earlier phase, we should still have to say that, much more than a million years after his departure from the chimpanzee level, man had merely advanced far enough to chip stone implements; because we find no other trace whatever of intelligence than this until near the close of the Palaeolithic period. If there is any mystery, it is in the slowness of man's development.

Let us further recollect that it is a common occurrence in the calendar of life for a particular organ to be especially developed in one member of a particular group more than in the others. The trunk of the elephant, the neck of the giraffe, the limbs of the horse or deer, the canines of the satire-toothed tiger, the wings of the bat, the colouring of the tiger, the horns of the deer, are so many examples in the mammal world alone. The brain is a useful organ like any other, and it is easy to conceive that the circumstances of one group may select it just as the environment of another group may lead to the selection of speed, weapons, or colouring. In fact, as we saw, there was so great and general an evolution of brain in the Tertiary Era that our modern mammals quite commonly have many times the brain of their Tertiary ancestors. Can we suggest any reasons why brain should be especially developed in the apes, and more particularly still in the ancestors of man?

The Primate group generally is a race of tree-climbers. The appearance of fruit on early Tertiary trees and the multiplication of carnivores explain this. The Primate is, except in a few robust cases, a particularly defenceless animal. When its earliest ancestors came in contact with fruit and nut-bearing trees, they developed climbing power and other means of defence and offense were sacrificed. Keenness of scent and range of hearing would now be of less moment, but sight would be stimulated, especially when soft-footed climbing carnivores came on the scene. There is, however, a much deeper significance in the adoption of climbing, and we must borrow a page from the modern physiology of the brain to understand it.

The stress laid in the modern education of young children on the use of the hands is not merely due to a feeling that they should handle objects as well as read about them. It is partly due to the belief of many distinguished physiologists that the training of the hands has a direct stimulating effect on the thought-centres in the brain. The centre in the cerebrum which controls the use of the hands is on the fringe of the region which seems to be concerned in mental operations. For reasons which will appear presently, we may add that the centres for controlling the muscles of the face and head are in the same region. Any finer training or the use of the hands will develop the centre for the fore limbs, and, on the principles, may react on the more important region of the cortex. Hence in turning the fore foot into a hand, for climbing and grasping purposes, the primitive Primate entered upon the path of brain-development. Even the earliest Primates show large brains in comparison with the small brains of their contemporaries.

It is a familiar fact in the animal world that when a certain group enters upon a particular path of evolution, some members of the group advance only a little way along it, some go farther, and some outstrip all the others. The development of social life among the bees will illustrate this. Hence we need not be puzzled by the fact that the lemurs have remained at one mental level, the monkeys at another, and the apes at a third. It is the common experience of life; and it is especially clear among the various races of men. A group becomes fitted to its environment, and, as long as its surroundings do not change, it does not advance. A related group, in a different environment, receives a particular stimulation, and advances. If, moreover, a group remains unstimulated for ages, it may become so rigid in its type that it loses the capacity to advance. It is generally believed that the lowest races of men, and even some of the higher races like the Australian aboriginals, are in this condition. We may expect this "unteachability" in a far more stubborn degree in the anthropoid apes, which have been adapted to an unchanging environment for a million years.

All that we need further suppose is—and it is one of the commonest episodes in terrestrial life—that one branch of the Miocene anthropoids, which were spread over a large part of the earth, received some stimulus to change which its cousins did not experience. It is sometimes suggested that social life was the great advantage which led to the superior development of mind in man. But such evidence as there is would lead us to suppose that primitive man was solitary, not social. The anthropoid apes are not social, but live in families, and are very unprogressive. On the other hand, the earliest remains of prehistoric man give no indication of social life. Fire-places, workshops, caves, etc., enter the story in a later phase. Some authorities on prehistoric man hold very strongly that during the greater part of the Old Stone Age (two-thirds, at least, of the human period) man wandered only in the company of his mate and children. [*]

* The point will be more fully discussed later. This account of prehistoric life is well seen in Mortillet's Prehistorique (1900). The lowest races also have no tribal life, and Professor Westermarck is of opinion that early man was not social.

We seem to have the most plausible explanation of the divergence of man from his anthropoid cousins in the fact that he left the trees of his and their ancestors. This theory has the advantage of being a fact—for the Ape-Man race of Java has already left the trees—and providing a strong ground for brain-advance. A dozen reasons might be imagined for his quitting the trees—migration, for instance, to a region in which food was more abundant, and carnivores less formidable, on the ground-level—but we will be content with the fact that he did. Such a change would lead to a more consistent adoption of the upright attitude, which is partly found in the anthropoid apes, especially the gibbons. The fore limb would be no longer a support of the body; the hand would be used more for grasping; and the hand-centre in the brain would be proportionately stimulated. The adoption of the erect attitude would further lead to a special development of the muscles of the head and face, the centre for which is in the same important region in the cortex. There would also be a direct stimulation of the brain, as, having neither weapons nor speed, the animal would rely all the more on sight and mind. If we further suppose that this primitive being extended the range of his hunting, from insects and small or dead birds to small land-animals, the stimulation would be all the greater. In a word, the very fact of a change from the trees to the ground suggests a line of brain-development which may plausibly be conceived, in the course of a million years, to evolve an Ape-Man out of a man-like ape. And we are not introducing any imaginary factor in this view of human origins.

The problem of the evolution of man is often approached in a frame of mind not far removed from that of the educated, but inexpert, European who stands before the lowly figure of the chimpanzee, and wonders by what miracle the gulf between it and himself was bridged. That is to lay a superfluous strain on the imagination. The proper term of comparison is the lowest type of human being known to us, since the higher types of living men have confessedly evolved from the lower. But even the lowest type of existing or recent savage is not the lowest level of humanity. Whether or no the Tasmanian or the Yahgan is a primitive remnant of the Old Stone Age, we have a far lower depth in the Java race. What we have first to do is to explain the advance to that level, in the course of many hundreds of thousands of years: a period fully a hundred times as long as the whole history of civilisation. Time itself is no factor in evolution, but in this case it is a significant condition. It means that, on this view of the evolution of man, we are merely assuming that an advance in brain-development took place between the Miocene and the Pleistocene, not similar to, but immeasurably less than, the advance which we know to have been made in the last fifty thousand years. In point of fact, the most mysterious feature of the evolution of man was its slowness. We shall see that, to meet the facts, we must suppose man to have made little or no progress during most of this vast period, and then to have received some new stimulation to develop. What it was we have now to inquire.



CHAPTER XIX. MAN AND THE GREAT ICE-AGE

In discussing the development of plants and animals during the Tertiary Era we have already perceived the shadow of the approaching Ice-Age. We found that in the course of the Tertiary the types which were more sensitive to cold gradually receded southward, and before its close Europe, Asia, and North America presented a distinctly temperate aspect. This is but the penumbra of the eclipse. When we pass the limits of the Tertiary Era, and enter the Quaternary, the refrigeration steadily proceeds, and, from temperate, the aspect of much of Europe and North America becomes arctic. From six to eight million square miles of the northern hemisphere are buried under fields of snow and ice, and even in the southern regions smaller glacial sheets spread from the foot of the higher ranges of mountains.

It is unnecessary to-day to explain at any length the evidences by which geologists trace this enormous glaciation of the northern hemisphere. There are a few works still in circulation in which popular writers, relying on the obstinacy of a few older geologists, speak lightly of the "nightmare" of the Ice-Age. But the age has gone by in which it could seriously be suggested that the boulders strewn along the east of Scotland—fragments of rock whose home we must seek in Scandinavia—were brought by the vikings as ballast for their ships. Even the more serious controversy, whether the scratches and the boulders which we find on the face of Northern Europe and America were due to floating or land ice, is virtually settled. Several decades of research have detected the unmistakable signs of glacial action over this vast area of the northern hemisphere. Most of Europe north of the Thames and the Danube, nearly all Canada and a very large part of the United States, and a somewhat less expanse of Northern Asia, bear to this day the deep scars of the thick, moving ice-sheets. Exposed rock-surfaces are ground and scratched, beds of pebbles are twisted and contorted hollows are scooped out, and moraines—the rubbish-heaps of the glaciers—are found on every side. There is now not the least doubt that, where the great Deinosaurs had floundered in semi-tropical swamps, where the figs and magnolias had later flourished, where the most industrious and prosperous hives of men are found to-day, there was, in the Pleistocene period, a country to which no parallel can be found outside the polar circles to-day.

The great revolution begins with the gathering of snows on the mountains. The Alps and Pyrenees had now, we saw, reached their full stature, and the gathering snows on their summits began to glide down toward the plains in rivers of ice. The Apennines (and even the mountains of Corsica), the Balkans, Carpathians, Caucasus, and Ural Mountains, shone in similar mantles of ice and snow. The mountains of Wales, the north of England, Scotland, and Scandinavia had even heavier burdens, and, as the period advanced, their sluggish streams of ice poured slowly over the plains. The trees struggled against the increasing cold in the narrowing tracts of green; the animals died, migrated to the south, or put on arctic coats. At length the ice-sheets of Scandinavia met the spreading sheets from Scotland and Wales, and crept over Russia and Germany, and an almost continuous mantle, from which only a few large areas of arctic vegetation peeped out, was thrown over the greater part of Europe. Ten thousand feet thick where it left the hills of Norway and Sweden, several thousand feet thick even in Scotland, the ice-sheet that resulted from the fusion of the glaciers gradually thinned as it went south, and ended in an irregular fringe across Central Europe. The continent at that time stretched westward beyond the Hebrides and some two hundred miles beyond Ireland. The ice-front followed this curve, casting icebergs into the Atlantic, then probably advanced up what is now the Bristol Channel, and ran across England and Europe, in a broken line, from Bristol to Poland. South of this line there were smaller ice-fields round the higher mountains, north of it almost the whole country presented the appearance that we find in Greenland to-day.

In North America the glaciation was even more extensive. About four million square miles of the present temperate zone were buried under ice and snow. From Greenland, Labrador, and the higher Canadian mountains the glaciers poured south, until, in the east, the mass of ice penetrated as far as the valley of the Mississippi. The great lakes of North America are permanent memorials of its Ice-Age, and over more than half the country we trace the imprint and the relics of the sheet. South America, Australia, Tasmania, and New Zealand had their glaciated areas. North Asia was largely glaciated, but the range of the ice-sheet is not yet determined in that continent.

This summary statement will convey some idea of the extraordinary phase through which the earth passed in the early part of the present geological era. But it must be added that a singular circumstance prolonged the glacial regime in the northern hemisphere. Modern geologists speak rather of a series of successive ice-sheets than of one definite Ice-Age. Some, indeed, speak of a series of Ice-Ages, but we need not discuss the verbal question. It is now beyond question that the ice-sheet advanced and retreated several times during the Glacial Epoch. The American and some English geologists distinguished six ice-sheets, with five intermediate periods of more temperate climate. The German and many English and French geologists distinguish four sheets and three interglacial epochs. The exact number does not concern us, but the repeated spread of the ice is a point of some importance. The various sheets differed considerably in extent. The wide range of the ice which I have described represents the greatest extension of the glaciation, and probably corresponds to the second or third of the six advances in Dr. Geikie's (and the American) classification.

Before we consider the biological effect of this great of refrigeration of the globe, we must endeavour to understand the occurrence itself. Here we enter a world of controversy, but a few suggestions at least may be gathered from the large literature of the subject, which dispel much of the mystery of the Great Ice-Age.

It was at one time customary to look out beyond the earth itself for the ultimate causes of this glaciation. Imagine the sheet of ice, which now spreads widely round the North Pole, shifted to another position on the surface of the planet, and you have a simple explanation of the occurrence. In other words, if we suppose that the axis of the earth does not consistently point in one direction—that the great ball does not always present the same average angle in relation to the sun—the poles will not always be where they are at present, and the Pleistocene Ice-Age may represent a time when the north pole was in the latitude of North Europe and North America. This opinion had to be abandoned. We have no trace whatever of such a constant shifting of the polar regions as it supposes, and, especially, we have no trace that the warm zone correspondingly shifted in the Pleistocene.

A much more elaborate theory was advanced by Dr. Croll, and is still entertained by many. The path of the earth round the sun is not circular, but elliptical, and there are times when the gravitational pull of the other planets increases the eccentricity of the orbit. It was assumed that there are periods of great length, separated from each other by still longer periods, when this eccentricity of the orbit is greatly exaggerated. The effect would be to prolong the winter and shorten the summer of each hemisphere in turn. The total amount of heat received would not alter, but there would be a long winter with less heat per hour, and a short summer with more heat. The short summer would not suffice to melt the enormous winter accumulations of ice and snow, and an ice-age would result. To this theory, again, it is objected that we do not find the regular succession of ice-ages in the story of the earth which the theory demands, and that there is no evidence of an alternation of the ice between the northern and southern hemispheres.

More recent writers have appealed to the sun itself, and supposed that some prolonged veiling of its photosphere greatly reduced the amount of heat emitted by it. More recently still it has been suggested that an accumulation of cosmic or meteoric dust in our atmosphere, or between us and the sun, had, for a prolonged period, the effect of a colossal "fire-screen." Neither of these suppositions would explain the localisation of the ice. In any case we need not have recourse to purely speculative accidents in the world beyond until it is clear that there were no changes in the earth itself which afford some explanation.

This is by no means clear. Some writers appeal to changes in the ocean currents. It is certain that a change in the course of the cold and warm currents of the ocean to-day might cause very extensive changes of climate, but there seems to be some confusion of ideas in suggesting that this might have had an equal, or even greater, influence in former times. Our ocean currents differ so much in temperature because the earth is now divided into very pronounced zones of climate. These zones did not exist before the Pliocene period, and it is not at all clear that any redistribution of currents in earlier times could have had such remarkable consequences. The same difficulty applies to wind-currents.

On the other hand, we have already, in discussing the Permian glaciation, discovered two agencies which are very effective in lowering the temperature of the earth. One is the rise of the land; the other is the thinning of the atmosphere. These are closely related agencies, and we found them acting in conjunction to bring about the Permian Ice-Age. Do we find them at work in the Pleistocene?

It is not disputed that there was a very considerable upheaval of the land, especially in Europe and North America, at the end of the Tertiary Era. Every mountain chain advanced, and our Alps, Pyrenees, Himalaya, etc., attained, for the first time, their present, or an even greater elevation. The most critical geologists admit that Europe, as a whole, rose 4000 feet above its earlier level. Such an elevation would be bound to involve a great lowering of the temperature. The geniality of the Oligocene period was due, like that of the earlier warm periods, to the low-lying land and very extensive water-surface. These conditions were revolutionised before the end of the Tertiary. Great mountains towered into the snow-line, and vast areas were elevated which had formerly been sea or swamp.

This rise of the land involved a great decrease in the proportion of moisture in the atmosphere. The sea surface was enormously lessened, and the mountains would now condense the moisture into snow or cloud to a vastly greater extent than had ever been known before There would also be a more active circulation of the atmosphere, the moist warm winds rushing upward towards the colder elevations and parting with their vapour. As the proportion of moisture in the atmosphere lessened the surface-heat would escape more freely into space, the general temperature would fall, and the evaporation—or production of moisture would be checked, while the condensation would continue. The prolonging of such conditions during a geological period can be understood to have caused the accumulation of fields of snow and ice in the higher regions. It seems further probable that these conditions would lead to a very considerable formation of fog and cloud, and under this protecting canopy the glaciers would creep further down toward the plains.

We have then to consider the possibility of a reduction of the quantity of carbon-dioxide in the atmosphere The inexpert reader probably has a very exaggerated idea of the fall in temperature that would be required to give Europe an Ice-Age. If our average temperature fell about 5-8 degrees C. below the average temperature of our time it would suffice; and it is further calculated that if the quantity of carbon-dioxide in our atmosphere were reduced by half, we should have this required fall in temperature. So great a reduction would not be necessary in view of the other refrigerating agencies. Now it is quite certain that the proportion of carbon-dioxide was greatly reduced in the Pleistocene. The forests of the Tertiary Era would steadily reduce it, but the extensive upheaval of the land at its close would be even more important. The newly exposed surfaces would absorb great quantities of carbon. The ocean, also, as it became colder, would absorb larger and larger quantities of carbon-dioxide. Thus the Pleistocene atmosphere, gradually relieved of its vapours and carbon-dioxide, would no longer retain the heat at the surface. We may add that the growth of reflective surfaces—ice, snow, cloud, etc.—would further lessen the amount of heat received from the sun.

Here, then, we have a series of closely related causes and effects which would go far toward explaining, if they do not wholly suffice to explain, the general fall of the earth's temperature. The basic cause is the upheaval of the land—a fact which is beyond controversy, the other agencies are very plain and recognisable consequences of the upheaval. There are, however, many geologists who do not think this explanation adequate.

It is pointed out, in the first place, that the glaciation seems to have come long after the elevation. The difficulty does not seem to be insurmountable. The reduction of the atmospheric vapour would be a gradual process, beginning with the later part of the elevation and culminating long afterwards. The reduction of the carbon-dioxide would be even more gradual. It is impossible to say how long it would take these processes to reach a very effective stage, but it is equally impossible to show that the interval between the upheaval and the glaciation is greater than the theory demands.

It is also said that we cannot on these principles understand the repeated advance and retreat of the ice-sheet.

This objection, again, seems to fail. It is an established fact that the land sank very considerably during the Ice-Age, and has risen again since the ice disappeared. We find that the crust in places sank so low that an arctic ocean bathed the slopes of some of the Welsh mountains; and American geologists say that their land has risen in places from 2000 to 3000 feet (Chamberlin) since the burden of ice was lifted from it. Here we have the possibility of an explanation of the advances and retreats of the glaciers. The refrigerating agencies would proceed until an enormous burden of ice was laid on the land of the northern hemisphere. The land apparently sank under the burden, the ice and snow melted at the lower level and there was a temperate interglacial period. But the land, relieved of its burden, rose once more, the exposed surface absorbed further quantities of carbon, and a fresh period of refrigeration opened. This oscillation might continue until the two sets of opposing forces were adjusted, and the crust reached a condition of comparative stability.

Finally, and this is the more serious difficulty, it is said that we cannot in this way explain the localisation of the glacial sheets. Why should Europe and North America in particular suffer so markedly from a general thinning of the atmosphere? The simplest answer is to suggest that they especially shared the rise of the land. Geology is not in a position either to prove or disprove this, and it remains only a speculative interpretation of the fact We know at least that there was a great uprise of land in Europe and North America in the Pliocene and Pleistocene and may leave the precise determination of the point to a later age. At the same time other local causes are not excluded. There may have been a large extension of the area of atmospheric depression which we have in the region of Greenland to-day.

When we turn to the question of chronology we have the same acute difference of opinion as we have found in regard to all questions of geological time. It used to be urged, on astronomical grounds, that the Ice-Age began about 240,000 years ago, and ended about 60,000 years ago, but the astronomical theory is, as I said, generally abandoned. Geologists, on the other hand, find it difficult to give even approximate figures. Reviewing the various methods of calculation, Professor Chamberlin concludes that the time of the first spread of the ice-sheet is quite unknown, the second and greatest extension of the glaciation may have been between 300,000 and a million years ago, and the last ice-extension from 20,000 to 60,000 years ago; but he himself attaches "very little value" to the figures. The chief ice-age was some hundreds of thousands of years ago, that is all we can say with any confidence.

In dismissing the question of climate, however, we should note that a very serious problem remains unsolved. As far as present evidence goes we seem to be free to hold that the ice-ages which have at long intervals invaded the chronicle of the earth were due to rises of the land. Upheaval is the one constant and clearly recognisable feature associated with, or preceding, ice-ages. We saw this in the case of the Cambrian, Permian, Eocene, and Pleistocene periods of cold, and may add that there are traces of a rise of mountains before the glaciation of which we find traces in the middle of the Archaean Era. There are problems still to be solved in connection with each of these very important ages, but in the rise of the land and consequent thinning of the atmosphere we seem to have a general clue to their occurrence. Apart from these special periods of cold, however, we have seen that there has been, in recent geological times, a progressive cooling of the earth, which we have not explained. Winter seems now to be a permanent feature of the earth's life, and polar caps are another recent, and apparently permanent, acquisition. I find no plausible reason assigned for this.

The suggestion that the disk of the sun is appreciably smaller since Tertiary days is absurd; and the idea that the earth has only recently ceased to allow its internal heat to leak through the crust is hardly more plausible. The cause remains to be discovered.

We turn now to consider the effect of the great Ice-Age, and the relation of man to it. The Permian revolution, to which the Pleistocene Ice-Age comes nearest in importance, wrought such devastation that the overwhelming majority of living things perished. Do we find a similar destruction of life, and selection of higher types, after the Pleistocene perturbation? In particular, had it any appreciable effect upon the human species?

A full description of the effect of the great Ice-Age would occupy a volume. The modern landscape in Europe and North America was very largely carved and modelled by the ice-sheet and the floods that ensued upon its melting. Hills were rounded, valleys carved, lakes formed, gravels and soils distributed, as we find them to-day. In its vegetal aspect, also, as we saw, the modern landscape was determined by the Pleistocene revolution. A great scythe slowly passed over the land. When the ice and snow had ended, and the trees and flowers, crowded in the southern area, slowly spread once more over the virgin soil, it was only the temperate species that could pass the zone guarded by the Alps and the Pyrenees. On the Alps themselves the Pleistocene population still lingers, their successful adaptation to the cold now preventing them from descending to the plains.

The animal world in turn was winnowed by the Pleistocene episode. The hippopotamus, crocodile, turtle, flamingo, and other warm-loving animals were banished to the warm zone. The mammoth and the rhinoceros met the cold by developing woolly coats, but the disappearance of the ice, which had tempted them to this departure, seems to have ended their fitness. Other animals which became adapted to the cold—arctic bears, foxes, seals, etc.—have retreated north with the ice, as the sheet melted. For hundreds of thousands of years Europe and North America, with their alternating glacial and interglacial periods, witnessed extraordinary changes and minglings of their animal population. At one time the reindeer, the mammoth, and the glutton penetrate down to the Mediterranean, in the next phase the elephant and hippopotamus again advance nearly to Central Europe. It is impossible here to attempt to unravel these successive changes and migrations. Great numbers of species were destroyed, and at length, when the climatic condition of the earth reached a state of comparative stability, the surviving animals settled in the geographical regions in which we find them to-day.

The only question into which we may enter with any fullness is that of the relation of human development to this grave perturbation of the condition of the globe. The problem is sometimes wrongly conceived. The chief point to be determined is not whether man did or did not precede the Ice-Age. As it is the general belief that he was evolved in the Tertiary, it is clear that he existed in some part of the earth before the Ice-Age. Whether he had already penetrated as far north as Britain and Belgium is an interesting point, but not one of great importance. We may, therefore, refrain from discussing at any length those disputed crude stone implements (Eoliths) which, in the opinion of many, prove his presence in northern regions before the close of the Tertiary. We may also now disregard the remains of the Java Ape-Man. There are authorities, such as Deniker, who hold that even the latest research shows these remains to be Pliocene, but it is disputed. The Java race may be a surviving remnant of an earlier phase of human evolution.

The most interesting subject for inquiry is the fortune of our human and prehuman forerunners during the Pliocene and Pleistocene periods. It may seem that if we set aside the disputable evidence of the Eoliths and the Java remains we can say nothing whatever on this subject. In reality a fact of very great interest can be established. It can be shown that the progress made during this enormous lapse of time—at least a million years—was remarkably slow. Instead of supposing that some extraordinary evolution took place in that conveniently obscure past, to which we can find no parallel within known times, it is precisely the reverse. The advance that has taken place within the historical period is far greater, comparatively to the span of time, than that which took place in the past.

To make this interesting fact clearer we must attempt to measure the progress made in the Pliocene and Pleistocene. We may assume that the precursor of man had arrived at the anthropoid-ape level by the middle of the Miocene period. He is not at all likely to have been behind the anthropoid apes, and we saw that they were well developed in the mid-Tertiary. Now we have a good knowledge of man as he was in the later stage of the Ice-Age—at least a million years later—and may thus institute a useful comparison and form some idea of the advance made.

In the later stages of the Pleistocene a race of men lived in Europe of whom we have a number of skulls and skeletons, besides vast numbers of stone implements. It is usually known as the Neanderthal race, as the first skeleton was found, in 1856, at Neanderthal, near Dusseldorf. Further skeletons were found at Spy, in Belgium, and Krapina, in Croatia. A skull formerly found at Gibraltar is now assigned to the same race. In the last five years a jaw of the same (or an earlier) age has been found at Mauer, near Heidelberg, and several skeletons have been found in France (La Vezere and Chapelle-aux-Saints). From these, and a few earlier fragments, we have a confident knowledge of the features of this early human race.

The highest appreciation of the Neanderthal man—a somewhat flattering appreciation, as we shall see—is that he had reached the level of the Australian black of to-day. The massive frontal ridges over his eyes, the very low, retreating forehead, the throwing of the mass of the brain toward the back of the head, the outthrust of the teeth and jaws, and the complete absence (in some cases) or very slight development of the chin, combine to give the head what the leading authorities call a "bestial" or "simian" aspect. The frame is heavy, powerful, and of moderate height (usually from two to four inches over five feet). The thigh-bones are much more curved than in modern man. We cannot enter here into finer anatomical details, but all the features are consistent and indicate a stage in the evolution from ape-man to savage man.

One point only calls for closer inquiry. Until a year or two ago it was customary to state that in cranial capacity also—that is to say, in the volume of brain-matter that the skull might contain—the Neanderthal race was intermediate between the Ape-Man and modern man. We saw above that the cranial capacity of the highest ape is about 600 cubic centimetres, and that of the Ape-Man (variously given as 850 and 950) is about 900. It was then added that the capacity of the Neanderthal race was about 1200, and that of civilised man (on the average) 1600. This seemed to be an effective and convincing indication of evolution, but recent writers have seriously criticised it. Sir Edwin Ray Lankester, Professor Sollas, and Dr. Keith have claimed in recent publications that the brain of Neanderthal man was as large as, if not larger than, that of modern man. [*] Professor Sollas even observes that "the brain increases in volume as we go backward." This is, apparently, so serious a reversal of the familiar statement in regard to the evolution of man that we must consider it carefully.

*See especially an address by Professor Sollas in the Quarterly Journal of the Geological Society, Vol. LXVI. (1910).

Largeness of brain in an individual is no indication of intelligence, and smallness of brain no proof of low mentality. Some of the greatest thinkers, such as Aristotle and Leibnitz, had abnormally small heads. Further, the size of the brain is of no significance whatever except in strict relation to the size and weight of the body. Woman has five or six ounces less brain-matter than man, but in proportion to her average size and the weight of the vital tissue of her body (excluding fat) she has as respectable a brain as man. When, however, these allowances have been made, it has usually been considered that the average brain of a race is in proportion to its average intelligence. This is not strictly true. The rabbit has a larger proportion of brain to body than the elephant or horse, and the canary a larger proportion than the chimpanzee. Professor Sollas says that the average cranial capacity of the Eskimo is 1546 cubic centimetres, or nearly that assigned to the average Parisian.

Clearly the question is very complex, and some of these recent authorities conclude that the cranial capacity, or volume of the brain, has no relation to intelligence, and therefore the size of the Neanderthal skull neither confirms nor disturbs the theory of evolution. The wise man will suspend his judgment until the whole question has been fully reconsidered. But I would point out that some of the recent criticisms are exaggerated. The Gibraltar skull is estimated by Professor Sollas himself to have a capacity of about 1260; and his conclusion that it is an abnormal or feminine skull rests on no positive grounds. The Chapelle-aux-Saints skull ALONE is proved to have the high capacity of 1620; and it is as yet not much more than a supposition that the earlier skulls had been wrongly measured. But, further, the great French authority, M. Boule, who measured the capacity of the Chapelle-aux Saints skull, observes [*] that "the anomaly disappears" on careful study. He assures us that a modern skull of the same dimensions would have a capacity of 1800-1900 cubic centimetres, and warns us that we must take into account the robustness of the body of primitive man. He concludes that the real volume of the Neanderthal brain (in this highest known specimen) is "slight in comparison with the volume of the brain lodged in the large heads of to-day," and that the "bestial or ape-like characters" of the race are not neutralised by this gross measurement.

*See his article in Anthropologie, Vol. XX. (1909), p. 257. As Professor Sollas mainly relies on Boule, it is important to see that there is a very great difference between the two.

We must therefore hesitate to accept the statement that primitive man had as large a brain, if not a larger brain, than a modern race. The basis is slender, and the proportion of brain to body-tissue has not been taken into account. On the other hand, the remains of this early race are, Professor Sollas says, "obviously more brutal than existing men in all the other ascertainable characters by which they differ from them." Nor are we confined to precarious measurements of skulls. We have the remains of the culture of this early race, and in them we have a surer trace of its mental development.