A closer examination of these early reptiles may be postponed until we come to speak of the "age of reptiles." We shall see that it is probable that an even higher type of animal, the mammal, was born in the throes of the Permian revolution. But enough has been said in vindication of the phrase which stands at the head of this chapter; and to show how the great Primary age of terrestrial life came to a close. With its new inhabitants the earth enters upon a fresh phase, and thousands of its earlier animals and plants are sealed in their primordial tombs, to await the day when man will break the seals and put flesh once more on the petrified bones.



CHAPTER XI. THE MIDDLE AGES OF THE EARTH

The story of the earth from the beginning of the Cambrian period to the present day was long ago divided by geologists into four great eras. The periods we have already covered—the Cambrian, Ordovician, Silurian, Devonian, Carboniferous, and Permian—form the Primary or Palaeozoic Era, to which the earlier Archaean rocks were prefixed as a barren and less interesting introduction. The stretch of time on which we now enter, at the close of the Permian, is the Secondary or Mesozoic Era. It will be closed by a fresh upheaval of the earth and disturbance of life-conditions in the Chalk period, and followed by a Tertiary Era, in which the earth will approach its modern aspect. At its close there will be another series of upheavals, culminating in a great Ice-age, and the remaining stretch of the earth's story, in which we live, will form the Quaternary Era.

In point of duration these four eras differ enormously from each other. If the first be conceived as comprising sixteen million years—a very moderate estimate—the second will be found to cover less than eight million years, the third less than three million years, and the fourth, the Age of Man, much less than one million years; while the Archaean Age was probably as long as all these put together. But the division is rather based on certain gaps, or "unconformities," in the geological record; and, although the breaches are now partially filled, we saw that they correspond to certain profound and revolutionary disturbances in the face of the earth. We retain them, therefore, as convenient and logical divisions of the biological as well as the geological chronicle, and, instead of passing from one geological period to another, we may, for the rest of the story, take these three eras as wholes, and devote a few chapters to the chief advances made by living things in each era. The Mesozoic Era will be a protracted reaction between two revolutions: a period of low-lying land, great sea-invasions, and genial climate, between two upheavals of the earth. The Tertiary Era will represent a less sharply defined depression, with genial climate and luxuriant life, between two such upheavals.

The Mesozaic ("middle life") Era may very fitly be described as the Middle Ages of life on the earth. It by no means occupies a central position in the chronicle of life from the point of view of time or antiquity, just as the Middle Ages of Europe are by no means the centre of the chronicle of mankind, but its types of animals and plants are singularly transitional between the extinct ancient and the actual modern types. Life has been lifted to a higher level by the Permian revolution. Then, for some millions of years, the sterner process of selection relaxes, the warm bosom of the earth swarms again with a teeming and varied population, and a rich material is provided for the next great application of drastic selective agencies. To a poet it might seem that nature indulges each succeeding and imperfect type of living thing with a golden age before it is dismissed to make place for the higher.

The Mesozoic opens in the middle of the great revolution described in the last chapter. Its first section, the Triassic period, is at first a mere continuation of the Permian. A few hundred species of animals and hardy plants are scattered over a relatively bleak and inhospitable globe. Then the land begins to sink once more. The seas spread in great arms over the revelled continents, the plant world rejoices in the increasing warmth and moisture, and the animals increase in number and variety. We pass into the Jurassic period under conditions of great geniality. Warm seas are found as far north and south as our present polar regions, and the low-lying fertile lands are covered again with rich, if less gigantic, forests, in which hordes of stupendous animals find ample nourishment. The mammal and the bird are already on the stage, but their warm coats and warm blood offer no advantage in that perennial summer, and they await in obscurity the end of the golden age of the reptiles. At the end of the Jurassic the land begins to rise once more. The warm, shallow seas drain off into the deep oceans, and the moist, swampy lands are dried. The emergence continues throughout the Cretaceous (Chalk) period. Chains of vast mountains rise slowly into the air in many parts of the earth, and a new and comparatively rapid change in the vegetation—comparable to that at the close of the Carboniferous—announces the second great revolution. The Mesozoic closes with the dismissal of the great reptiles and the plants on which they fed, and the earth is prepared for its new monarchs, the flowering plants, the birds, and the mammals.

How far this repeated levelling of the land after its repeated upheavals is due to a real sinking of the crust we cannot as yet determine. The geologist of our time is disposed to restrict these mysterious rises and falls of the crust as much as possible. A much more obvious and intelligible agency has to be considered. The vast upheaval of nearly all parts of the land during the Permian period would naturally lead to a far more vigorous scouring of its surface by the rains and rivers. The higher the land, the more effectively it would be worn down. The cooler summits would condense the moisture, and the rains would sweep more energetically down the slopes of the elevated continents. There would thus be a natural process of levelling as long as the land stood out high above the water-line, but it seems probable that there was also a real sinking of the crust. Such subsidences have been known within historic times.

By the end of the Triassic—a period of at least two million years—the sea had reconquered a vast proportion of the territory wrested from it in the Permian revolution. Most of Europe, west of a line drawn from the tip of Norway to the Black Sea, was under water—generally open sea in the south and centre, and inland seas or lagoons in the west. The invasion of the sea continued, and reached its climax, in the Jurassic period. The greater part of Europe was converted into an archipelago. A small continent stood out in the Baltic region. Large areas remained above the sea-level in Austria, Germany, and France. Ireland, Wales, and much of Scotland were intact, and it is probable that a land bridge still connected the west of Europe with the east of America. Europe generally was a large cluster of islands and ridges, of various sizes, in a semi-tropical sea. Southern Asia was similarly revelled, and it is probable that the seas stretched, with little interruption, from the west of Europe to the Pacific. The southern continent had deep wedges of the sea driven into it. India, New Zealand, and Australia were successively detached from it, and by the end of the Mesozoic it was much as we find it to-day. The Arctic continent (north of Europe) was flooded, and there was a great interior sea in the western part of the North American continent.

This summary account of the levelling process which went on during the Triassic and Jurassic will prepare us to expect a return of warm climate and luxurious life, and this the record abundantly evinces. The enormous expansion of the sea—a great authority, Neumayr, believes that it was the greatest extension of the sea that is known in geology—and lowering of the land would of itself tend to produce this condition, and it may be that the very considerable volcanic activity, of which we find evidence in the Permian and Triassic, had discharged great volumes of carbon-dioxide into the atmosphere.

Whatever the causes were, the earth has returned to paradisiacal conditions. The vast ice-fields have gone, the scanty and scrubby vegetation is replaced by luscious forests of cycads, conifers, and ferns, and warmth-loving animals penetrate to what are now the Arctic and Antarctic regions. Greenland and Spitzbergen are fragments of a continent that then bore a luxuriant growth of ferns and cycads, and housed large reptiles that could not now live thousands of miles south of it. England, and a large part of Europe, was a tranquil blue coral-ocean, the fringes of its islands girt with reefs such as we find now only three thousand miles further south, with vast shoals of Ammonites, sometimes of gigantic size, preying upon its living population or evading its monstrous sharks; while the sunlit lands were covered with graceful, palmlike cycads and early yews and pines and cypresses, and quaint forms of reptiles throve on the warm earth or in the ample swamps, or rushed on outstretched wings through the purer air.

It was an evergreen world, a world, apparently, of perpetual summer. No trace is found until the next period of an alternation of summer and winter—no trees that shed their leaves annually, or show annual rings of growth in the wood—and there is little trace of zones of climate as yet. It is true that the sensitive Ammonites differ in the northern and the southern latitudes, but, as Professor Chamberlin says, it is not clear that the difference points to a diversity of climate. We may conclude that the absence of corals higher than the north of England implies a more temperate climate further north, but what Sir A. Geikie calls (with slight exaggeration) "the almost tropical aspect" of Greenland warns us to be cautious. The climate of the mid-Jurassic was very much warmer and more uniform than the climate of the earth to-day. It was an age of great vital expansion. And into this luxuriant world we shall presently find a fresh period of elevation, disturbance, and cold breaking with momentous evolutionary results. Meantime, we may take a closer look at these interesting inhabitants of the Middle Ages of the earth, before they pass away or are driven, in shrunken regiments, into the shelter of the narrowing tropics.

The principal change in the aspect of the earth, as the cold, arid plains and slopes of the Triassic slowly yield the moist and warm ow-lying lands of the Jurassic, to consists in the character of the vegetation. It is wholly intermediate in its forms between that of the primitive forests and that of the modern world. The great Cryptogams of the Carboniferous world—the giant Club-mosses and their kindred—have been slain by the long period of cold and drought. Smaller Horsetails (sometimes of a great size, but generally of the modern type) and Club-mosses remain, but are not a conspicuous feature in the landscape. On the other hand, there is as yet—apart from the Conifers—no trace of the familiar trees and flowers and grasses of the later world. The vast majority of the plants are of the cycad type. These—now confined to tropical and subtropical regions—with the surviving ferns, the new Conifers, and certain trees of the ginkgo type, form the characteristic Mesozoic vegetation.

A few words in the language of the modern botanist will show how this vegetation harmonises with the story of evolution. Plants are broadly divided into the lower kingdom of the Cryptogams (spore-bearing) and the upper kingdom of the Phanerogams (seed-bearing). As we saw, the Primary Era was predominantly the age of Cryptogams; the later periods witness the rise and supremacy of the Phanerogams. But these in turn are broadly divided into a less advanced group, the Gymnosperms, and a more advanced group, the Angiosperms or flowering plants. And, just as the Primary Era is the age of Cryptogams, the Secondary is the age of Gymnosperms, and the Tertiary (and present) is the age of Angiosperms. Of about 180,000 species of plants in nature to-day more than 100,000 are Angiosperms; yet up to the end of the Jurassic not a single true Angiosperm is found in the geological record.

This is a broad manifestation of evolution, but it is not quite an accurate statement, and its inexactness still more strongly confirms the theory of evolution. Though the Primary Era was predominantly the age of Cryptogams, we saw that a very large number of seed-bearing plants, with very mixed characters, appeared before its close. It thus prepares the way for the cycads and conifers and ginkgoes of the Mesozoic, which we may conceive as evolved from one or other branch of the mixed Carboniferous vegetation. We next find that the Mesozoic is by no means purely an age of Gymnosperms. I do not mean merely that the Angiosperms appear in force before its close, and were probably evolved much earlier. The fact is that the Gymnosperms of the Mesozoic are often of a curiously mixed character, and well illustrate the transition to the Angiosperms, though they may not be their actual ancestors. This will be clearer if we glance in succession at the various types of plant which adorned and enriched the Jurassic world.

The European or American landscape—indeed, the aspect of the earth generally, for there are no pronounced zones of climate—is still utterly different from any that we know to-day. No grass carpets the plains; none of the flowers or trees with which we are familiar, except conifers, are found in any region. Ferns grow in great abundance, and have now reached many of the forms with which we are acquainted. Thickets of bracken spread over the plains; clumps of Royal ferns and Hartstongues spring up in moister parts. The trees are conifers, cycads, and trees akin to the ginkgo, or Maidenhair Tree, of modern Japan. Cypresses, yews, firs, and araucarias (the Monkey Puzzle group) grow everywhere, though the species are more primitive than those of today. The broad, fan-like leaves and plum-like fruit of the ginkgoales, of which the temple-gardens of Japan have religiously preserved a solitary descendant, are found in the most distant regions. But the most frequent and characteristic tree of the Jurassic landscape is the cycad.

The cycads—the botanist would say Cycadophyta or Cycadales, to mark them off from the cycads of modern times—formed a third of the whole Jurassic vegetation, while to-day they number only about a hundred species in 180,000, and are confined to warm latitudes. All over the earth, from the Arctic to the Antarctic, their palm-like foliage showered from the top of their generally short stems in the Jurassic. But the most interesting point about them is that a very large branch of them (the Bennettiteae) went far beyond the modern Gymnosperm in their flowers and fruit, and approached the Angiosperms. Their fructifications "rivalled the largest flowers of the present day in structure and modelling" (Scott), and possibly already gave spots of sober colour to the monotonous primitive landscape. On the other hand, they approached the ferns so much more closely than modern cycads do that it is often impossible to say whether Jurassic remains must be classed as ferns or cycads.

We have here, therefore, a most interesting evolutionary group. The botanist finds even more difficulty than the zoologist in drawing up the pedigrees of his plants, but the general features of the larger groups which he finds in succession in the chronicle of the earth point very decisively to evolution. The seed-bearing ferns of the Coal-forest point upward to the later stage, and downward to a common origin with the ordinary spore-bearing ferns. Some of them are "altogether of a cycadean type" (Scott) in respect of the seed. On the other hand, the Bennettiteae of the Jurassic have the mixed characters of ferns, cycads, and flowering plants, and thus, in their turn, point downward to a lower ancestry and upward to the next great stage in plant-development. It is not suggested that the seed-ferns we know evolved into the cycads we know, and these in turn into our flowering plants. It is enough for the student of evolution to see in them so many stages in the evolution of plants up to the Angiosperm level. The gaps between the various groups are less rigid than scientific men used to think.

Taller than the cycads, firmer in the structure of the wood, and destined to survive in thousands of species when the cycads would be reduced to a hundred, were the pines and yews and other conifers of the Jurassic landscape. We saw them first appearing, in the stunted Walchias and Voltzias, during the severe conditions of the Permian period. Like the birds and mammals they await the coming of a fresh period of cold to give them a decided superiority over the cycads. Botanists look for their ancestors in some form related to the Cordaites of the Coal-forest. The ginkgo trees seem to be even more closely related to the Cordaites, and evolved from an early and generalised branch of that group. The Cordaites, we may recall, more or less united in one tree the characters of the conifer (in their wood) and the cycad (in their fruit).

So much for the evolutionary aspect of the Jurassic vegetation in itself. Slender as the connecting links are, it points clearly enough to a selection of higher types during the Permian revolution from the varied mass of the Carboniferous flora, and it offers in turn a singularly varied and rich group from which a fresh selection may choose yet higher types. We turn now to consider the animal population which, directly or indirectly, fed upon it, and grew with its growth. To the reptiles, the birds, and the mammals, we must devote special chapters. Here we may briefly survey the less conspicuous animals of the Mesozoic Epoch.

The insects would be one of the chief classes to benefit by the renewed luxuriance of the vegetation. The Hymenopters (butterflies) have not yet appeared. They will, naturally, come with the flowers in the next great phase of organic life. But all the other orders of insects are represented, and many of our modern genera are fully evolved. The giant insects of the Coal-forest, with their mixed patriarchal features, have given place to more definite types. Swarms of dragon-flies, may-flies, termites (with wings), crickets, and cockroaches, may be gathered from the preserved remains. The beetles (Coleopters) have come on the scene in the Triassic, and prospered exceedingly. In some strata three-fourths of the insects are beetles, and as we find that many of them are wood-eaters, we are not surprised. Flies (Dipters) and ants (Hymenopters) also are found, and, although it is useless to expect to find the intermediate forms of such frail creatures, the record is of some evolutionary interest. The ants are all winged. Apparently there is as yet none of the remarkable division of labour which we find in the ants to-day, and we may trust that some later period of change may throw light on its origin.

Just as the growth of the forests—for the Mesozoic vegetation has formed immense coal-beds in many parts of the world, even in Yorkshire and Scotland—explains this great development of the insects, they would in their turn supply a rich diet to the smaller land animals and flying animals of the time. We shall see this presently. Let us first glance at the advances among the inhabitants of the seas.

The most important and stimulating event in the seas is the arrival of the Ammonite. One branch of the early shell-fish, it will be remembered, retained the head of its naked ancestor, and lived at the open mouth of its shell, thus giving birth to the Cephalopods. The first form was a long, straight, tapering shell, sometimes several feet long. In the course of time new forms with curved shells appeared, and began to displace the straight-shelled. Then Cephalopods with close-coiled shells, like the nautilus, came, and—such a shell being an obvious advantage—displaced the curved shells. In the Permian, we saw, a new and more advanced type of the coiled-shell animal, the Ammonite, made its appearance, and in the Triassic and Jurassic it becomes the ogre or tyrant of the invertebrate world. Sometimes an inch or less in diameter, it often attained a width of three feet or more across the shell, at the aperture of which would be a monstrous and voracious mouth.

The Ammonites are not merely interesting as extinct monsters of the earth's Middle Ages, and stimulating terrors of the deep to the animals on which they fed. They have an especial interest for the evolutionist. The successive chambers which the animal adds, as it grows, to the habitation of its youth, leave the earlier chambers intact. By removing them in succession in the adult form we find an illustration of the evolution of the elaborate shell of the Jurassic Ammonite. It is an admirable testimony to the validity of the embryonic law we have often quoted—that the young animal is apt to reproduce the past stages of its ancestry—that the order of the building of the shell in the late Ammonite corresponds to the order we trace in its development in the geological chronicle. About a thousand species of Ammonites were developed in the Mesozoic, and none survived the Mesozoic. Like the Trilobites of the Primary Era, like the contemporary great reptiles on land, the Ammonites were an abortive growth, enjoying their hour of supremacy until sterner conditions bade them depart. The pretty nautilus is the only survivor to-day of the vast Mesozoic population of coiled-shell Cephalopods.

A rival to the Ammonite appeared in the Triassic seas, a formidable forerunner of the cuttle-fish type of Cephalopod. The animal now boldly discards the protecting and confining shell, or spreads over the outside of it, and becomes a "shell-fish" with the shell inside. The octopus of our own time has advanced still further, and become the most powerful of the invertebrates. The Belemnite, as the Mesozoic cuttle-fish is called, attained so large a size that the internal bone, or pen (the part generally preserved), is sometimes two feet in length. The ink-bags of the Belemnite also are sometimes preserved, and we see how it could balk a pursuer by darkening the waters. It was a compensating advantage for the loss of the shell.

In all the other classes of aquatic animals we find corresponding advances. In the remaining Molluscs the higher or more effective types are displacing the older. It is interesting to note that the oyster is fully developed, and has a very large kindred, in the Mesozoic seas. Among the Brachiopods the higher sloping-shoulder type displaces the square-shoulder shells. In the Crustacea the Trilobites and Eurypterids have entirely disappeared; prawns and lobsters abound, and the earliest crab makes its appearance in the English Jurassic rocks. This sudden arrival of a short-tailed Crustacean surprises us less when we learn that the crab has a long tail in its embryonic form, but the actual line of its descent is not clear. Among the Echinoderms we find that the Cystids and Blastoids have gone, and the sea-lilies reach their climax in beauty and organisation, to dwindle and almost disappear in the last part of the Mesozoic. One Jurassic sea-lily was found to have 600,000 distinct ossicles in its petrified frame. The free-moving Echinoderms are now in the ascendant, the sea-urchins being especially abundant. The Corals are, as we saw, extremely abundant, and a higher type (the Hexacoralla) is superseding the earlier and lower (Tetracoralla).

Finally, we find a continuous and conspicuous advance among the fishes. At the close of the Triassic and during the Jurassic they seem to undergo profound and comparatively rapid changes. The reason will, perhaps, be apparent in the next chapter, when we describe the gigantic reptiles which feed on them in the lakes and shore-waters. A greater terror than the shark had appeared in their environment. The Ganoids and Dipneusts dwindle, and give birth to their few modern representatives. The sharks with crushing teeth diminish in number, and the sharp-toothed modern shark attains the supremacy in its class, and evolves into forms far more terrible than any that we know to-day. Skates and rays of a more or less modern type, and ancestral gar-pikes and sturgeons, enter the arena. But the most interesting new departure is the first appearance, in the Jurassic, of bony-framed fishes (Teleosts). Their superiority in organisation soon makes itself felt, and they enter upon the rapid evolution which will, by the next period, give them the first place in the fish world.

Over the whole Mesozoic world, therefore, we find advance and the promise of greater advance. The Permian stress has selected the fittest types to survive from the older order; the Jurassic luxuriance is permitting a fresh and varied expansion of life, in preparation for the next great annihilation of the less fit and selection of the more fit. Life pauses before another leap. The Mesozoic earth—to apply to it the phrase which a geologist has given to its opening phase—welcomes the coming and speeds the parting guest. In the depths of the ocean a new movement is preparing, but we have yet to study the highest forms of Mesozoic life before we come to the Cretaceous disturbances.



CHAPTER XII. THE AGE OF REPTILES

From one point of view the advance of life on the earth seems to proceed not with the even flow of a river, but in the successive waves of an oncoming tide. It is true that we have detected a continuous advance behind all these rising and receding waves, yet their occurrence is a fact of some interest, and not a little speculation has been expended on it. When the great procession of life first emerges out of the darkness of Archaean times, it deploys into a spreading world of strange Crustaceans, and we have the Age of Trilobites. Later there is the Age of Fishes, then of Cryptogams and Amphibia, and then of Cycads and Reptiles, and there will afterwards be an Age of Birds and Mammals, and finally an Age of Man. But there is no ground for mystic speculation on this circumstance of a group of organisms fording the earth for a few million years, and then perishing or dwindling into insignificance. We shall see that a very plain and substantial process put an end to the Age of the Cycads, Ammonites, and Reptiles, and we have seen how the earlier dynasties ended.

The phrase, however, the Age of Reptiles, is a fitting and true description of the greater part of the Mesozoic Era, which lies, like a fertile valley, between the Permian and the Chalk upheavals. From the bleak heights of the Permian period, or—more probably—from its more sheltered regions, in which they have lingered with the ferns and cycads, the reptiles spread out over the earth, as the summer of the Triassic period advances. In the full warmth and luxuriance of the Jurassic they become the most singular and powerful army that ever trod the earth. They include small lizard-like creatures and monsters more than a hundred feet in length. They swim like whales in the shallow seas; they shrink into the shell of the giant turtle; they rear themselves on towering hind limbs, like colossal kangaroos; they even rise into the air, and fill it with the dragons of the fairy tale. They spread over the whole earth from Australia to the Arctic circle. Then the earth seems to grow impatient of their dominance, and they shrink towards the south, and struggle in a diminished territory. The colossal monsters and the formidable dragons go the way of all primitive life, and a ragged regiment of crocodiles, turtles, and serpents in the tropics, with a swarm of smaller creatures in the fringes of the warm zone, is all that remains, by the Tertiary Era, of the world-conquering army of the Mesozoic reptiles.

They had appeared, as we said, in the Permian period. Probably they had been developed during the later Carboniferous, since we find them already branched into three orders, with many sub-orders, in the Permian. The stimulating and selecting disturbances which culminated in the Permian revolution had begun in the Carboniferous. Their origin is not clear, as the intermediate forms between them and the amphibia are not found. This is not surprising, if we may suppose that some of the amphibia had, in the growing struggle, pushed inland, or that, as the land rose and the waters were drained in certain regions, they had gradually adopted a purely terrestrial life, as some of the frogs have since done. In the absence of water their frames would not be preserved and fossilised. We can, therefore, understand the gap in the record between the amphibia and the reptiles. From their structure we gather that they sprang from at least two different branches of the amphibia. Their remains fall into two great groups, which are known as the Diapsid and the Synapsid reptiles. The former seem to be more closely related to the Microsauria, or small salamander-like amphibia of the Coal-forest; the latter are nearer to the Labyrinthodonts. It is not suggested that these were their actual ancestors, but that they came from the same early amphibian root.

We find both these groups, in patriarchal forms, in Europe, North America, and South Africa during the Permian period. They are usually moderate in size, but in places they seem to have found good conditions and prospered. A few years ago a Permian bed in Russia yielded a most interesting series of remains of Synapsid reptiles. Some of them were large vegetarian animals, more than twelve feet in length; others were carnivores with very powerful heads and teeth as formidable as those of the tiger. Another branch of the same order lived on the southern continent, Gondwana Land, and has left numerous remains in South Africa. We shall see that they are connected by many authorities with the origin of the mammals. [*] The other branch, the Diapsids, are represented to-day by the curiously primitive lizard of New Zealand, the tuatara (Sphenodon, or Hatteria), of which I have seen specimens, nearly two feet in length, that one did not care to approach too closely. The Diapsids are chiefly interesting, however, as the reputed ancestors of the colossal reptiles of the Jurassic age and the birds.

* These Synapsid reptiles are more commonly known as Pareiasauria or Theromorpha.

The purified air of the Permian world favoured the reptiles' being lung-breathers, but the cold would check their expansion for a time. The reptile, it is important to remember' usually leaves its eggs to be hatched by the natural warmth of the ground. But as the cold of the Permian yielded to a genial climate and rich vegetation in the course of the Triassic, the reptiles entered upon their memorable development. The amphibia were now definitely ousted from their position of dominance. The increase of the waters had at first favoured them, and we find more than twenty genera, and some very large individuals, of the amphibia in the Triassic. One of them, the Mastodonsaurus, had a head three feet long and two feet wide. But the spread of the reptiles checked them, and they shrank rapidly into the poor and defenceless tribe which we find them in nature to-day.

To follow the prolific expansion of the reptiles in the semi-tropical conditions of the Jurassic age is a task that even the highest authorities approach with great diffidence. Science is not yet wholly agreed in the classification of the vast numbers of remains which the Mesozoic rocks have yielded, and the affinities of the various groups are very uncertain. We cannot be content, however, merely to throw on the screen, as it were, a few of the more quaint and monstrous types out of the teeming Mesozoic population, and describe their proportions and peculiarities. They fall into natural and intelligible groups or orders, and their features are closely related to the differing regions of the Jurassic world. While, therefore, we must abstain from drawing up settled genealogical trees, we may, as we review in succession the monsters of the land, the waters, and the air, glance at the most recent and substantial conjectures of scientific men as to their origin and connections.

The Deinosaurs (or "terrible reptiles"), the monarchs of the land and the swamps, are the central and outstanding family of the Mesozoic reptiles. As the name implies, this group includes most of the colossal animals, such as the Diplodocus, which the illustrated magazine has made familiar to most people. Fortunately the assiduous research of American geologists and their great skill and patience in restoring the dead forms enable us to form a very fair picture of this family of medieval giants and its remarkable ramifications. [*]

* See, besides the usual authorities, a valuable paper by Dr. R. S. Lull, "Dinosaurian Distribution" (1910).

The Diapsid reptiles of the Permian had evolved a group with horny, parrot-like beaks, the Rhyncocephalia (or "beak-headed" reptiles), of which the tuatara of New Zealand is a lingering representative. New Zealand seems to have been cut off from the southern continent at the close of the Permian or beginning of the Triassic, and so preserved for us that very interesting relic of Permian life. From some primitive level of this group, it is generally believed, the great Deinosaurs arose. Two different orders seem to have arisen independently, or diverged rapidly from each other, in different parts of the world. One group seems to have evolved on the "lost Atlantis," the land between Western Europe and America, whence they spread westward to America, eastward over Europe, and southward to the continent which still united Africa and Australia. We find their remains in all these regions. Another stock is believed to have arisen in America.

Both these groups seem to have been more or less biped, rearing themselves on large and powerful hind limbs, and (in some cases, at least) probably using their small front limbs to hold or grasp their food. The first group was carnivorous, the second herbivorous; and, as the reptiles of the first group had four or five toes on each foot, they are known as the Theropods (or "beast-footed" ), while those of the second order, which had three toes, are called the Ornithopods (or "bird-footed"). Each of them then gave birth to an order of quadrupeds. In the spreading waters and rich swamps of the later Triassic some of the Theropods were attracted to return to an amphibious life, and became the vast, sprawling, ponderous Sauropods, the giants in a world of giants. On the other hand, a branch of the vegetarian Ornithopods developed heavy armour, for defence against the carnivores, and became, under the burden of its weight, the quadrupedal and monstrous Stegosauria and Ceratopsia. Taking this instructive general view of the spread of the Deinosaurs as the best interpretation of the material we have, we may now glance at each of the orders in succession.

The Theropods varied considerably in size and agility. The Compsognathus was a small, active, rabbit-like creature, standing about two feet high on its hind limbs, while the Megalosaurs stretched to a length of thirty feet, and had huge jaws armed with rows of formidable teeth. The Ceratosaur, a seventeen-foot-long reptile, had hollow bones, and we find this combination of lightness and strength in several members of the group. In many respects the group points more or less significantly toward the birds. The brain is relatively large, the neck long, and the fore limbs might be used for grasping, but had apparently ceased to serve as legs. Many of the Theropods were evidently leaping reptiles, like colossal kangaroos, twenty or more feet in length when they were erect. It is the general belief that the bird began its career as a leaping reptile, and the feathers, or expanded scales, on the front limbs helped at first to increase the leap. Some recent authorities hold, however, that the ancestor of the bird was an arboreal reptile.

To the order of the Sauropods belong most of the monsters whose discovery has attracted general attention in recent years. Feeding on vegetal matter in the luscious swamps, and having their vast bulk lightened by their aquatic life, they soon attained the most formidable proportions. The admirer of the enormous skeleton of Diplodocus (which ran to eighty feet) in the British Museum must wonder how even such massive limbs could sustain the mountain of flesh that must have covered those bones. It probably did not walk so firmly as the skeleton suggests, but sprawled in the swamps or swam like a hippopotamus. But the Diplodocus is neither the largest nor heaviest of its family. The Brontosaur, though only sixty feet long, probably weighed twenty tons. We have its footprints in the rocks to-day, each impression measuring about a square yard. Generally, it is the huge thigh-bones of these monsters that have survived, and give us an idea of their size. The largest living elephant has a femur scarcely four feet long, but the femur of the Atlantosaur measures more than seventy inches, and the femur of the Brachiosaur more than eighty. Many of these Deinosaurs must have measured more than a hundred feet from the tip of the snout to the end of the tail, and stood about thirty feet high from the ground. The European Sauropods did not, apparently, reach the size of their American cousins—so early did the inferiority of Europe begin—but our Ceteosaur seems to have been about fifty feet long and ten feet in height. Its thigh-bone was sixty-four inches long and twenty-seven inches in circumference at the shaft. And in this order of reptiles, it must be remembered, the bones are solid.

To complete the picture of the Sauropods, we must add that the whole class is characterised by the extraordinary smallness of the brain. The twenty-ton Brontosaur had a brain no larger than that of a new-born human infant. Quite commonly the brain of one of these enormous animals is no larger than a man's fist. It is true that, as far as the muscular and sexual labour was concerned, the brain was supplemented by a great enlargement of the spinal cord in the sacral region (at the top of the thighs). This inferior "brain" was from ten to twenty times as large as the brain in the skull. It would, however, be fully occupied with the movement of the monstrous limbs and tail, and the sex-life, and does not add in the least to the "mental" power of the Sauropods. They were stupid, sluggish, unwieldy creatures, swollen parasites upon a luxuriant vegetation, and we shall easily understand their disappearance at the end of the Mesozoic Era, when the age of brawn will yield to an age of brain.

The next order of the Deinosaurs is that of the biped vegetarians, the Ornithopods, which gradually became heavily armoured and quadrupedal. The familiar Iguanodon is the chief representative of this order in Europe. Walking on its three-toed hind limbs, its head would be fourteen or fifteen feet from the ground. The front part of its jaws was toothless and covered with horn. It had, in fact, a kind of beak, and it also approached the primitive bird in the structure of its pelvis and in having five toes on its small front limbs. Some of the Ornithopods, such as the Laosaur, were small (three or four feet in height) and active, but many of the American specimens attained a great size. The Camptosaur, which was closely related to the Iguanodon in structure, was thirty feet from the snout to the end of the tail, and the head probably stood eighteen feet from the ground. One of the last great representatives of the group in America, the Trachodon, about thirty feet in length, had a most extraordinary head. It was about three and a half feet in length, and had no less than 2000 teeth lining the mouth cavity. It is conjectured that it fed on vegetation containing a large proportion of silica.

In the course of the Jurassic, as we saw, a branch of these biped, bird-footed vegetarians developed heavy armour, and returned to the quadrupedal habit. We find them both in Europe and America, and must suppose that the highway across the North Atlantic still existed.

The Stegosaur is one of the most singular and most familiar representatives of the group in the Jurassic. It ran to a length of thirty feet, and had a row of bony plates, from two to three feet in height, standing up vertically along the ridge of its back, while its tail was armed with formidable spikes. The Scleidosaur, an earlier and smaller (twelve-foot) specimen, also had spines and bony plates to protect it. The Polacanthus and Ankylosaur developed a most effective armour-plating over the rear. As we regard their powerful armour, we seem to see the fierce-toothed Theropods springing from the rear upon the poor-mouthed vegetarians. The carnivores selected the vegetarians, and fitted them to survive. Before the end of the Mesozoic, in fact, the Ornithopods became aggressive as well as armoured. The Triceratops had not only an enormous skull with a great ridged collar round the neck, but a sharp beak, a stout horn on the nose, and two large and sharp horns on the top of the head. We will see something later of the development of horns. The skulls of members of the Ceratops family sometimes measured eight feet from the snout to the ridge of the collar. They were, however, sluggish and stupid monsters, with smaller brains even than the Sauropods.

Such, in broad outline, was the singular and powerful family of the Mesozoic Deinosaurs. Further geological research in all parts of the world will, no doubt, increase our knowledge of them, until we can fully understand them as a great family throwing out special branches to meet the different conditions of the crowded Jurassic age. Even now they afford a most interesting page in the story of evolution, and their total disappearance from the face of the earth in the next geological period will not be unintelligible. We turn from them to the remaining orders of the Jurassic reptiles.

In the popular mind, perhaps, the Ichthyosaur and Plesiosaur are the typical representatives of that extinct race. The two animals, however, belong to very different branches of the reptile world, and are by no means the most formidable of the Mesozoic reptiles. Many orders of the land reptiles sent a branch into the waters in an age which, we saw, was predominantly one of water-surface. The Ichthyosauria ("fish-reptiles") and Thalattosauria ("sea-reptiles") invaded the waters at their first expansion in the later Triassic. The latter groups soon became extinct, but the former continued for some millions of years, and became remarkably adapted to marine life, like the whale at a later period.

The Ichthyosaur of the Jurassic is a remarkably fish-like animal. Its long tapering frame—sometimes forty feet in length, but generally less than half that length—ends in a dip of the vertebral column and an expansion of the flesh into a strong tail-fin. The terminal bones of the limbs depart more and more from the quadruped type, until at last they are merely rows of circular bony plates embedded in the broad paddle into which the limb has been converted. The head is drawn out, sometimes to a length of five feet, and the long narrow jaws are set with two formidable rows of teeth; one specimen has about two hundred teeth. In some genera the teeth degenerate in the course of time, but this merely indicates a change of diet. One fossilised Ichthyosaur of the weaker-toothed variety has been found with the remains of two hundred Belemnites in its stomach. It is a flash of light on the fierce struggle and carnage which some recent writers have vainly striven to attenuate. The eyes, again, which may in the larger animals be fifteen inches in diameter, are protected by a circle of radiating bony plates. In fine, the discovery of young developed skeletons inside the adult frames has taught us that the Ichthgosaur had become viviparous, like the mammal. Cutting its last connection with the land, on which it originated it ceased to lay eggs, and developed the young within its body.

The Ichthyosaur came of the reptile group which we have called the Diapsids. The Plesiosaur seems to belong to the Synapsid branch. In the earlier Mesozoic we find partially aquatic representatives of the line, like the Nothosaur, and in the later Plesiosaur the adaptation to a marine life is complete. The skin has lost its scales, and the front limbs are developed into powerful paddles, sometimes six feet in length. The neck is drawn out until, in some specimens, it is found to consist of seventy-six vertebrae: the longest neck in the animal world. It is now doubted, however, if the neck was very flexible, and, as the jaws were imperfectly joined, the common picture of the Plesiosaur darting its snake-like neck in all directions to seize its prey is probably wrong. It seems to have lived on small food, and been itself a rich diet to the larger carnivores. We find it in all the seas of the Mesozoic world, varying in length from six to forty feet, but it is one of the sluggish and unwieldy forms that are destined to perish in the coming crisis.

The last, and perhaps the most interesting, of the doomed monsters of the Mesozoic was the Pterosaur, or "flying reptile." It is not surprising that in the fierce struggle which is reflected in the arms and armour of the great reptiles, a branch of the family escaped into the upper region. We have seen that there were leaping reptiles with hollow bones, and although the intermediate forms are missing, there is little doubt that the Pterosaur developed from one or more of these leaping Deinosaurs. As it is at first small, when it appears in the early Jurassic—it is disputed in the late Triassic—it probably came from a small and agile Deinosaur, hunted by the carnivores, which relied on its leaping powers for escape. A flapperlike broadening of the fore limbs would help to lengthen the leap, and we must suppose that this membrane increased until the animal could sail through the air, like the flying-fish, and eventually sustain its weight in the air. The wing is, of course, not a feathery frame, as in the bird, but a special skin spreading between the fore limb and the side of the body. In the bat this skin is supported by four elongated fingers of the hand, but in the Pterosaur the fifth (or fourth) finger alone—which is enormously elongated and strengthened—forms its outer frame. It is as if, in flying experiments, a man were to have a web of silk stretching from his arm and an extension of his little finger to the side of his body.

From the small early specimens in the early Jurassic the flying reptiles grow larger and larger until the time of their extinction in the stresses of the Chalk upheaval. Small Pterosaurs continue throughout the period, but from these bat-like creatures we rise until we come to such dragons as the American Pteranodon, with a stretch of twenty-two feet between its extended wings and jaws about four feet long. There were long-tailed Pterosaurs (Ramphorhyncus), sometimes with a rudder-like expansion of the end of the tail, and short-tailed Pterosaurs (Pterodactyl), with compact bodies and keeled breasts, like the bird. In the earlier part of the period they all have the heavy jaws and numerous teeth of the reptile, with four or five well-developed fingers on the front limbs. In the course of time they lose the teeth—an advantage in the distribution of the weight of the body while flying—and develop horny beaks. In the gradual shaping of the breast-bone and head, also, they illustrate the evolution of the bird-form.

But the birds were meantime developing from a quite different stock, and would replace the Pterosaurs at the first change in the environment. There is ground for thinking that these flying reptiles were warm-blooded like the birds. Their hollow bones seem to point to the effective breathing of a warm-blooded animal, and the great vitality they would need in flying points toward the same conclusion. Their brain, too, approached that of the bird, and was much superior to that of the other reptiles. But they had no warm coats to retain their heat, no clavicle to give strength to the wing machinery, and, especially in the later period, they became very weak in the hind limbs (and therefore weak or slow in starting their flight). The coming selection will therefore dismiss them from the scene, with the Deinosaurs and Ammonites, and retain the better organised bird as the lord of the air.

There remain one or two groups of the Mesozoic reptiles which are still represented in nature. The turtle-group (Chelonia) makes its appearance in the Triassic and thrives in the Jurassic. Its members are extinct and primitive forms of the thick-shelled reptiles, but true turtles, both of marine and fresh water, abound before the close of the Mesozoic. The sea-turtles attain an enormous size. Archelon, one of the primitive types, measured about twelve feet across the shell. Another was thirteen feet long and fifteen feet from one outstretched flipper to the other. In the Chalk period they form more than a third of the reptile remains in some regions. They are extremely interesting in that they show, to some extent, the evolution of their characteristic shell. In some of the larger specimens the ribs have not yet entirely coalesced.

The Crocodilians also appear in the later Triassic, abound in the Jurassic, and give way before the later types, the true Crocodiles, in the Cretaceous. They were marine animals with naked skin, a head and neck something like that of the Ichthyosaur, and paddles like those of the Plesiosaur. Their back limbs, however, were not much changed after their adaptation to life in the sea, and it is concluded that they visited the land to lay their eggs. The Teleosaur was a formidable narrow-spouted reptile, somewhat resembling the crocodiles of the Ganges in the external form of the jaws. The modern crocodiles, which replaced this ancient race of sea-crocodiles, have a great advantage over them in the fact that their nostrils open into the mouth in its lower depths. They can therefore close their teeth on their prey under water and breathe through the nose.

Snakes are not found until the close of the Mesozoic, and do not figure in its characteristic reptile population. We will consider them later. But there was a large group of reptiles in the later Mesozoic seas which more or less correspond to the legendary idea of a sea-serpent. These Dolichosaurs ("long reptiles") appear at the beginning of the Chalk period, and develop into a group, the Mososaurians, which must have added considerably to the terrors of the shore-waters. Their slender scale-covered bodies were commonly twenty to thirty feet in length. The supreme representative of the order, the Mososaur, of which about forty species are known, was sometimes seventy-five feet long. It had two pairs of paddles—so that the name of sea-serpent is very imperfectly applicable—and four rows of formidable teeth on the roof of its mouth. Like the Deinosaurs and Pterosaurs, the order was doomed to be entirely extinguished after a brief supremacy in its environment.

From this short and summary catalogue the reader will be able to form some conception of the living inhabitants of the Mesozoic world. It is assuredly the Age of Reptiles. Worms, snails, and spiders were, we may assume, abundant enough, and a great variety of insects flitted from tree to tree or sheltered in the fern brakes. But the characteristic life, in water and on land, was the vast and diversified family of the reptiles. In the western and the eastern continent, and along the narrowing bridge that still united them, in the northern hemisphere and the southern, and along every ridge of land that connected them, these sluggish but formidable monsters filled the stage. Every conceivable device in the way of arms and armour, brute strength and means of escape, seemed to be adopted in their development, as if they were the final and indestructible outcome of the life-principle. And within a single geological period the overwhelming majority of them, especially the larger and more formidable of them, were ruthlessly slain, leaving not a single descendant on the earth. Let us see what types of animals were thus preferred to them in the next great application of selective processes.



CHAPTER XIII. THE BIRD AND THE MAMMAL

In one of his finest stories, Sur La Pierre Blanche, Anatole France has imagined a group of Roman patricians discussing the future of their Empire. The Christians, who are about to rise to power on their ruin, they dismiss with amiable indifference as one of the little passing eccentricities of the religious life of their time. They have not the dimmest prevision, even as the dream of a possibility, that in a century or two the Empire of Rome will lie in the dust, and the cross will tower above all its cities from York to Jerusalem. If we might for a moment endow the animals of the Mesozoic world with AEsopian wisdom, we could imagine some such discussion taking place between a group of Deinosaur patricians. They would reflect with pride on the unshakable empire of the reptiles, and perhaps glance with disdain at two types of animals which hid in the recesses or fled to the hills of the Jurassic world. And before another era of the earth's story opened, the reptile race would be dethroned, and these hunted and despised and feeble eccentricities of Mesozoic life would become the masters of the globe.

These two types of organisms were the bird and the mammal. Both existed in the Jurassic, and the mammals at least had many representatives in the Triassic. In other words, they existed, with all their higher organisation, during several million years without attaining power. The mammals remained, during at least 3,000,000 years, a small and obscure caste, immensely overshadowed by the small-brained reptiles. The birds, while making more progress, apparently, than the mammals, were far outnumbered by the flying reptiles until the last part of the Mesozoic. Then there was another momentous turn of the wheel of fate, and they emerged from their obscurity to assume the lordship of the globe.

In earlier years, when some serious hesitation was felt by many to accept the new doctrine of evolution, a grave difficulty was found in the circumstance that new types—not merely new species and new genera, but new orders and even sub-classes—appeared in the geological record quite suddenly. Was it not a singular coincidence that in ALL cases the intermediate organisms between one type and another should have wholly escaped preservation? The difficulty was generally due to an imperfect acquaintance with the conditions of the problem. The fossil population of a period is only that fraction of its living population which happened to be buried in a certain kind of deposit under water of a certain depth. We shall read later of insects being preserved in resin (amber), and we have animals (and even bacteria) preserved in trees from the Coal-forests. Generally speaking, however, the earth has buried only a very minute fraction of its land-population. Moreover, only a fraction of the earth's cemeteries have yet been opened. When we further reflect that the new type of organism, when it first appears, is a small and local group, we see what the chances are of our finding specimens of it in a few scattered pages of a very fragmentary record of the earth's life. We shall see that we have discovered only about ten skeletons or fragments of skeletons of the men who lived on the earth before the Neolithic period; a stretch of some hundreds of thousands of years, recorded in the upper strata of the earth.

Whatever serious difficulty there ever was in this scantiness of intermediate types is amply met by the fact that every fresh decade of search in the geological tombs brings some to light. We have seen many instances of this—the seed-bearing ferns and flower-bearing cycads, for example, found in the last decade—and will see others. But one of the most remarkable cases of the kind now claims our attention. The bird was probably evolved in the late Triassic or early Jurassic. It appears in abundance, divided into several genera, in the Chalk period. Luckily, two bird-skeletons have been found in the intermediate period, the Jurassic, and they are of the intermediate type, between the reptile and the bird, which the theory of evolution would suggest. But for the fortunate accident of these two birds being embedded in an ancient Bavarian mud-layer, which happened to be opened, for commercial purposes, in the second half of the nineteenth century, critics of evolution—if there still were any in the world of science—might be repeating to-day that the transition from the reptile to the bird was unthinkable in theory and unproven in fact.

The features of the Archaeopteryx ("primitive bird") have been described so often, and such excellent pictorial restorations of its appearance may now be seen, that we may deal with it briefly. We have in it a most instructive combination of the characters of the bird and the reptile. The feathers alone, the imprint of which is excellently preserved in the fine limestone, would indicate its bird nature, but other anatomical distinctions are clearly seen in it. "There is," says Dr. Woodward, "a typical bird's 'merrythought' between the wings, and the hind leg is exactly that of a perching bird." In other words, it has the shoulder-girdle and four-toed foot, as well as the feathers, of a bird. On the other hand, it has a long tail (instead of a terminal tuft of feathers as in the bird) consisting of twenty-one vertebrae, with the feathers springing in pairs from either side; it has biconcave vertebrae, like the fishes, amphibia, and reptiles; it has teeth in its jaws; and it has three complete fingers, free and clawed, on its front limbs.

As in the living Peripatus, therefore, we have here a very valuable connecting link between two very different types of organisms. It is clear that one of the smaller reptiles—the Archaeopteryx is between a pigeon and a crow in size—of the Triassic period was the ancestor of the birds. Its most conspicuous distinction was that it developed a coat of feathers. A more important difference between the bird and the reptile is that the heart of the bird is completely divided into four chambers, but, as we saw, this probably occurred also in the other flying reptiles. It may be said to be almost a condition of the greater energy of a flying animal. When the heart has four complete chambers, the carbonised blood from the tissues of the body can be conveyed direct to the lungs for purification, and the aerated blood taken direct to the tissues, without any mingling of the two. In the mud-fish and amphibian, we saw, the heart has two chambers (auricles) above, but one (ventricle) below, in which the pure and impure blood mingle. In the reptiles a partition begins to form in the lower chamber. In the turtle it is so nearly complete that the venous and the arterial blood are fairly separated; in the crocodile it is quite complete, though the arteries are imperfectly arranged. Thus the four-chambered heart of the bird and mammal is not a sudden and inexplicable development. Its advantage is enormous in a cold climate. The purer supply of blood increases the combustion in the tissues, and the animal maintains its temperature and vitality when the surrounding air falls in temperature. It ceases to be "cold-blooded."

But the bird secures a further advantage, and here it outstrips the flying reptile. The naked skin of the Pterosaur would allow the heat to escape so freely when the atmosphere cooled that a great strain would be laid on its vitality. A man lessens the demand on his vitality in cold regions by wearing clothing. The bird somehow obtained clothing, in the shape of a coat of feathers, and had more vitality to spare for life-purposes in a falling temperature. The reptile is strictly limited to one region, the bird can pass from region to region as food becomes scarce.

The question of the origin of the feathers can be discussed only from the speculative point of view, as they are fully developed in the Archaeopteryx, and there is no approach toward them in any other living or fossil organism. But a long discussion of the problem has convinced scientific men that the feathers are evolved from the scales of the reptile ancestor. The analogy between the shedding of the coat in a snake and the moulting of a bird is not uninstructive. In both cases the outer skin or epidermis is shedding an old growth, to be replaced by a new one. The covering or horny part of the scale and the feather are alike growths from the epidermis, and the initial stages of the growth have certain analogies. But beyond this general conviction that the feather is a development of the scale, we cannot proceed with any confidence. Nor need we linger in attempting to trace the gradual modification of the skeleton, owing to the material change in habits. The horny beak and the reduction of the toes are features we have already encountered in the reptile, and the modification of the pelvis, breast-bone, and clavicle are a natural outcome of flight.

In the Chalk period we find a large number of bird remains, of about thirty different species, and in some respects they resume the story of the evolution of the bird. They are widely removed from our modern types of birds, and still have teeth in the jaws. They are of two leading types, of which the Ichthyornis and Hesperornis are the standard specimens. The Ichthyornis was a small, tern-like bird with the power of flight strongly developed, as we may gather from the frame of its wings and the keel-shaped structure of its breast-bone. Its legs and feet were small and slender, and its long, slender jaws had about twenty teeth on each side at the bottom. No modern bird has teeth; though the fact that in some modern species we find the teeth appearing in a rudimentary form is another illustration of the law that animals tend to reproduce ancestral features in their development. A more reptilian character in the Ichthyornis group is the fact that, unlike any modern bird, but like their reptile ancestors, they had biconcave vertebrae. The brain was relatively poor. We are still dealing with a type intermediate in some respects between the reptile and the modern bird. The gannets, cormorants, and pelicans are believed to descend from some branch of this group.

The other group of Cretaceous birds, of the Hesperornis type, show an actual degeneration of the power of flight through adaptation to an environment in which it was not needed, as happened, later, in the kiwi of New Zealand, and is happening in the case of the barn-yard fowl. These birds had become divers. Their wings had shrunk into an abortive bone, while their powerful legs had been peculiarly fitted for diving. They stood out at right angles to the body, and seem to have developed paddles. The whole frame suggests that the bird could neither walk nor fly, but was an excellent diver and swimmer. Not infrequently as large as an ostrich (five to six feet high), with teeth set in grooves in its jaws, and the jaws themselves joined as in the snake, with a great capacity of bolting its prey, the Hesperornis would become an important element in the life of the fishes. The wing-fingers have gone, and the tail is much shortened, but the grooved teeth and loosely jointed jaws still point back to a reptilian ancestry.

These are the only remains of bird-life that we find in the Mesozoic rocks. Admirably as they illustrate the evolution of the bird from the reptile, they seem to represent a relatively poor development and spread of one of the most advanced organisms of the time. It must be understood that, as we shall see, the latter part of the Chalk period does not belong to the depression, the age of genial climate, which I call the Middle Ages of the earth, but to the revolutionary period which closes it. We may say that the bird, for all its advances in organisation, remains obscure and unprosperous as long as the Age of Reptiles lasts. It awaits the next massive uplift of the land and lowering of temperature.

In an earlier chapter I hinted that the bird and the mammal may have been the supreme outcomes of the series of disturbances which closed the Primary Epoch and devastated its primitive population. As far as the bird is concerned, this may be doubted on the ground that it first appears in the upper or later Jurassic, and is even then still largely reptilian in character. We must remember, however, that the elevation of the land and the cold climate lasted until the second part of the Triassic, and it is generally agreed that the bird may have been evolved in the Triassic. Its slow progress after that date is not difficult to understand. The advantage of a four-chambered heart and warm coat would be greatly reduced when the climate became warmer. The stimulus to advance would relax. The change from a coat of scales to a coat of feathers obviously means adaptation to a low temperature, and there is nothing to prevent us from locating it in the Triassic, and indeed no later known period of cold in which to place it.

It is much clearer that the mammals were a product of the Permian revolution. They not only abound throughout the Jurassic, in which they are distributed in more than thirty genera, but they may be traced into the Triassic itself. Both in North America and Europe we find the teeth and fragments of the jaws of small animals which are generally recognised as mammals. We cannot, of course, from a few bones deduce that there already, in the Triassic, existed an animal with a fully developed coat of fur and an apparatus, however crude, in the breast for suckling the young. But these bones so closely resemble the bones of the lowest mammals of to-day that this seems highly probable. In the latter part of the long period of cold it seems that some reptile exchanged its scales for tufts of hair, developed a four-chambered heart, and began the practice of nourishing the young from its own blood which would give the mammals so great an ascendancy in a colder world.

Nor can we complain of any lack of evidence connecting the mammal with a reptile ancestor. The earliest remains we find are of such a nature that the highest authorities are still at variance as to whether they should be classed as reptilian or mammalian. A skull and a fore limb from the Triassic of South Africa (Tritylodon and Theriodesmus) are in this predicament. It will be remembered that we divided the primitive reptiles of the Permian period into two great groups, the Diapsids and Synapsids (or Theromorphs). The former group have spread into the great reptiles of the Jurassic; the latter have remained in comparative obscurity. One branch of these Theromorph reptiles approach the mammals so closely in the formation of the teeth that they have received the name "of the Theriodonts", or "beast-toothed" reptiles. Their teeth are, like those of the mammals, divided into incisors, canines (sometimes several inches long), and molars; and the molars have in some cases developed cusps or tubercles. As the earlier remains of mammals which we find are generally teeth and jaws, the resemblance of the two groups leads to some confusion in classifying them, but from our point of view it is not unwelcome. It narrows the supposed gulf between the reptile and the mammal, and suggests very forcibly the particular branch of the reptiles to which we may look for the ancestry of the mammals. We cannot say that these Theriodont reptiles were the ancestors of the mammals. But we may conclude with some confidence that they bring us near to the point of origin, and probably had at least a common ancestor with the mammals.

The distribution of the Theriodonts suggests a further idea of interest in regard to the origin of the mammals. It would be improper to press this view in the present state of our knowledge, yet it offers a plausible theory of the origin of the mammals. The Theriodonts seem to have been generally confined to the southern continent, Gondwana Land (Brazil to Australia), of which an area survives in South Africa. It is there also that we find the early disputed remains of mammals. Now we saw that, during the Permian, Gondwana Land was heavily coated with ice, and it seems natural to suppose that the severe cold which the glacial fields would give to the whole southern continent was the great agency in the evolution of the highest type of the animal world. From this southern land the new-born mammals spread northward and eastward with great rapidity. Fitted as they were to withstand the rigorous conditions which held the reptiles and amphibia in check, they seemed destined to attain at once the domination of the earth. Then, as we saw, the land was revelled once more until its surface broke into a fresh semi-tropical luxuriance, and the Deinosaurs advanced to their triumph. The mammals shrank into a meagre and insignificant population, a scattered tribe of small insect-eating animals, awaiting a fresh refrigeration of the globe.

The remains of these interesting early mammals, restricted, as they generally are, to jaws and teeth and a few other bones that cannot in themselves be too confidently distinguished from those of certain reptiles, may seem insufficient to enable us to form a picture of their living forms. In this, however, we receive a singular and fortunate assistance. Some of them are found living in nature to-day, and their distinctly reptilian features would, even if no fossil remains were in existence, convince us of the evolution of the mammals.

The southern continent on which we suppose the mammals to have originated had its eastern termination in Australia. New Zealand seems to have been detached early in the Mesozoic, and was never reached by the mammals. Tasmania was still part of the Australian continent. To this extreme east of the southern continent the early mammals spread, and then, during either the Jurassic or the Cretaceous, the sea completed its inroad, and severed Australia permanently from the rest of the earth. The obvious result of this was to shelter the primitive life of Australia from invasion by higher types, especially from the great carnivorous mammals which would presently develop. Australia became, in other words, a "protected area," in which primitive types of life were preserved from destruction, and were at the same time sheltered from those stimulating agencies which compelled the rest of the world to advance. "Advance Australia" is the fitting motto of the present human inhabitants of that promising country; but the standard of progress has been set up in a land which had remained during millions of years the Chinese Empire of the living world. Australia is a fragment of the Middle Ages of the earth, a province fenced round by nature at least three million years ago and preserving, amongst its many invaluable types of life, representatives of that primitive mammal population which we are seeking to understand.

It is now well known that the Duckbill or Platypus (Ornithorhyncus) and the Spiny Anteater (Echidna) of Australia and Tasmania—with one representative of the latter in New Guinea, which seems to have been still connected—are semi-reptilian survivors of the first animals to suckle their young. Like the reptiles they lay tough-coated eggs and have a single outlet for the excreta, and they have a reptilian arrangement of the bones of the shoulder-girdle; like the mammals, they have a coat of hair and a four-chambered heart, and they suckle the young. Even in their mammalian features they are, as the careful research of Australian zoologists has shown, of a transitional type. They are warm-blooded, but their temperature is much lower than that of other mammals, and varies appreciably with the temperature of their surroundings. [*] Their apparatus for suckling the young is primitive. There are no teats, and the milk is forced by the mother through simple channels upon the breast, from which it is licked by the young. The Anteater develops her eggs in a pouch. They illustrate a very early stage in the development of a mammal from a reptile; and one is almost tempted to see in their timorous burrowing habits a reminiscence of the impotence of the early mammals after their premature appearance in the Triassic.

* See Lucas and Le Soulf's Animals of Australia, 1909.

The next level of mammal life, the highest level that it attains in Australia (apart from recent invasions), is the Marsupial. The pouched animals (kangaroo, wallaby, etc.) are the princes of pre-human life in Australia, and represent the highest point that life had reached when that continent was cut off from the rest of the world. A few words on the real significance of the pouch, from which they derive their name, will suffice to explain their position in the story of evolution.

Among the reptiles the task of the mother ends, as a rule, with the laying of the egg. One or two modern reptiles hatch the eggs, or show some concern for them, but the characteristic of the reptile is to discharge its eggs upon the warm earth and trouble no further about its young. It is a reminiscence of the warm primitive earth. The bird and mammal, born of the cooling of the earth, exhibit the beginning of that link between mother and offspring which will prove so important an element in the higher and later life of the globe. The bird assists the development of the eggs with the heat of her own body, and feeds the young. The mammal develops the young within the body, and then feeds them at the breast.

But there is a gradual advance in this process. The Duckbill lays its eggs just like the reptile, but provides a warm nest for them at the bottom of its burrow. The Anteater develops a temporary pouch in its body, when it lays an egg, and hatches the egg in it. The Marsupial retains the egg in its womb until the young is advanced in development, then transfers the young to the pouch, and forces milk into its mouth from its breasts. The real reason for this is that the Marsupial falls far short of the higher mammals in the structure of the womb, and cannot fully develop its young therein. It has no placenta, or arrangement by which the blood-vessels of the mother are brought into connection with the blood-vessels of the foetus, in order to supply it with food until it is fully developed. The Marsupial, in fact, only rises above the reptile in hatching the egg within its own body, and then suckling the young at the breast.

These primitive mammals help us to reconstruct the mammal life of the Mesozoic Epoch. The bones that we have are variously described in geological manuals as the remains of Monotremes, Marsupials, and Insectivores. Many of them, if not most, were no doubt insect-eating animals, but there is no ground for supposing that what are technically known as Insectivores (moles and shrews) existed in the Mesozoic. On the other hand, the lower jaw of the Marsupial is characterised by a peculiar hooklike process, and this is commonly found in Mesozoic jaws. This circumstance, and the witness of Australia, permit us, perhaps, to regard the Jurassic mammals as predominantly marsupial. It is more difficult to identify Monotreme remains, but the fact that Monotremes have survived to this day in Australia, and the resemblance of some of the Mesozoic teeth to those found for a time in the young Duckbill justify us in assuming that a part of the Mesozoic mammals correspond to the modern Monotremes. Not single specimen of any higher, or placental, mammal has yet been found in the whole Mesozoic Era.

We must, however, beware of simply transferring to the Mesozoic world the kinds of Monotremes and Marsupials which we know in nature to-day. In some of the excellent "restorations" of Mesozoic life which are found in recent illustrated literature the early mammal is represented with an external appearance like that of the Duckbill. This is an error, as the Duckbill has been greatly modified in its extremities and mouth-parts by its aquatic and burrowing habits. As we have no complete skeletons of these early mammals we must abstain from picturing their external appearance. It is enough that the living Monotreme and Marsupial so finely illustrate the transition from a reptilian to a mammalian form. There may have been types more primitive than the Duckbill, and others between the Duckbill and the Marsupial. It seems clear, at least, that two main branches, the Monotremes and Marsupials, arose from the primitive mammalian root. Whether either of these became in turn the parent of the higher mammals we will inquire later. We must first consider the fresh series of terrestrial disturbances which, like some gigantic sieve, weeded out the grosser types of organisms, and cleared the earth for a rapid and remarkable expansion of these primitive birds and mammals.

We have attended only to a few prominent characters in tracing the line of evolution, but it will be understood that an advance in many organs of the body is implied in these changes. In the lower mammals the diaphragm, or complete partition between the organs of the breast and those of the abdomen, is developed. It is not a sudden and mysterious growth, and its development in the embryo to-day corresponds to the suggestion of its development which the zoologist gathers from the animal series. The ear also is now fully developed. How far the fish has a sense of hearing is not yet fully determined, but the amphibian certainly has an organ for the perception of waves of sound. Parts of the discarded gill-arches are gradually transformed into the three bones of the mammal's internal ear; just as other parts are converted into mouth cartilages, and as—it is believed—one of the gill clefts is converted into the Eustachian tube. In the Monotreme and Marsupial the ear-hole begins to be covered with a shell of cartilage; we have the beginning of the external ear. The jaws, which are first developed in the fish, now articulate more perfectly with the skull. Fat-glands appear in the skin, and it is probably from a group of these that the milk-glands are developed. The origin of the hairs is somewhat obscure. They are not thought to be, like the bird's feathers, modifications of the reptile's scales, but to have been evolved from other structures in the skin, possibly under the protection of the scales.

My purpose is, however, rather to indicate the general causes of the onward advance of life than to study organs in detail—a vast subject—or construct pedigrees. We therefore pass on to consider the next great stride that is taken by the advancing life of the earth. Millions of years of genial climate and rich vegetation have filled the earth with a prolific and enormously varied population. Over this population the hand of natural selection is outstretched, as it were, and we are about to witness another gigantic removal of older types of life and promotion of those which contain the germs of further advance. As we have already explained, natural selection is by no means inactive during these intervening periods of warmth. We have seen the ammonites and reptiles, and even the birds and mammals, evolve into hundreds of species during the Jurassic period. The constant evolution of more effective types of carnivores and their spread into new regions, the continuous changes in the distribution of land and water, the struggle for food in a growing population, and a dozen other causes, are ever at work. But the great and comprehensive changes in the face of the earth which close the eras of the geologist seem to give a deeper and quicker stimulus to its population and result in periods of especially rapid evolution. Such a change now closes the Mesozoic Era, and inaugurates the age of flowering plants, of birds, and of mammals.



CHAPTER XIV. IN THE DAYS OF THE CHALK

In accordance with the view of the later story of the earth which was expressed on an earlier page, we now come to the second of the three great revolutions which have quickened the pulse of life on the earth. Many men of science resent the use of the word revolution, and it is not without some danger. It was once thought that the earth was really shaken at times by vast and sudden cataclysms, which destroyed its entire living population, so that new kingdoms of plants and animals had to be created. But we have interpreted the word revolution in a very different sense. The series of changes and disturbances to which we give the name extended over a period of hundreds of thousands of years, and they were themselves, in some sense, the creators of new types of organisms. Yet they are periods that stand out peculiarly in the comparatively even chronicle of the earth. The Permian period transformed the face of the earth; it lifted the low-lying land into a massive relief, drew mantles of ice over millions of miles of its surface, set volcanoes belching out fire and fumes in many parts, stripped it of its great forests, and slew the overwhelming majority of its animals. On the scale of geological time it may be called a revolution.

It must be confessed that the series of disturbances which close the Secondary and inaugurate the Tertiary Era cannot so conveniently be summed up in a single formula. They begin long before the end of the Mesozoic, and they continue far into the Tertiary, with intervals of ease and tranquillity. There seems to have been no culminating point in the series when the uplifted earth shivered in a mantle of ice and snow. Yet I propose to retain for this period—beginning early in the Cretaceous (Chalk) period and extending into the Tertiary—the name of the Cretaceous Revolution. I drew a fanciful parallel between the three revolutions which have quickened the earth since the sluggish days of the Coal-forest and the three revolutionary movements which have changed the life of modern Europe. It will be remembered that, whereas the first of these European revolutions was a sharp and massive upheaval, the second consisted in a more scattered and irregular series of disturbances, spread over the fourth and fifth decades of the nineteenth century; but they amounted, in effect, to a revolution.

So it is with the Cretaceous Revolution. In effect it corresponds very closely to the Permian Revolution. On the physical side it includes a very considerable rise of the land over the greater part of the globe, and the formation of lofty chains of mountains; on the botanical side it means the reduction of the rich Mesozoic flora to a relatively insignificant population, and the appearance and triumphant spread of the flowering plants, on the zoological side it witnesses the complete extinction of the Ammonites, Deinosaurs, and Pterosaurs, an immense reduction of the reptile world generally, and a victorious expansion of the higher insects, birds, and mammals; on the climatic side it provides the first definite evidence of cold zones of the earth and cold seasons of the year, and seems to represent a long, if irregular, period of comparative cold. Except, to some extent, the last of these points, there is no difference of opinion, and therefore, from the evolutionary point of view, the Cretaceous period merits the title of a revolution. All these things were done before the Tertiary period opened.

Let us first consider the fundamental and physical aspect of this revolution, the upheaval of the land. It began about the close of the Jurassic period. Western and Central Europe emerged considerably from the warm Jurassic sea, which lay on it and had converted it into an archipelago. In North-western America also there was an emergence of large areas of land, and the Sierra and Cascade ranges of mountains were formed about the same time. For reasons which will appear later we must note carefully this rise of land at the very beginning of the Cretaceous period.

However, the sea recovered its lost territory, or compensation for it, and the middle of the Cretaceous period witnessed a very considerable extension of the waters over America, Europe, and southern Asia. The thick familiar beds of chalk, which stretch irregularly from Ireland to the Crimea, and from the south of Sweden to the south of France, plainly tell of an overlying sea. As is well known, the chalk consists mainly of the shells or outer frames of minute one-celled creatures (Thalamophores) which float in the ocean, and form a deep ooze at its bottom with their discarded skeletons. What depth this ocean must have been is disputed, and hardly concerns us. It is clear that it must have taken an enormous period for microscopic shells to form the thick masses of chalk which cover so much of southern and eastern England. On the lowest estimates the Cretaceous period, which includes the deposit of other strata besides chalk, lasted about three million years. And as people like to have some idea of the time since these things happened, I may add that, on the lowest estimate (which most geologists would at least double), it is about three million years since the last stretches of the chalk-ocean disappeared from the surface of Europe.

But while our chalk cliffs conjure up a vision of England lying deep—at least twenty or thirty fathoms deep—below a warm ocean, in which gigantic Ammonites and Belemnites and sharks ply their deadly trade, they also remind us of the last phase of the remarkable life of the earth's Middle Ages. In the latter part of the Cretaceous the land rises. The chalk ocean of Europe is gradually reduced to a series of inland seas, separated by masses and ridges of land, and finally to a series of lakes of brackish water. The masses of the Pyrenees and Alps begin to rise; though it will not be until a much later date that they reach anything like their present elevation. In America the change is even greater. A vast ridge rises along the whole western front of the continent, lifting and draining it, from Alaska to Cape Horn. It is the beginning of the Rocky Mountains and the Andes. Even during the Cretaceous period there had been rich forests of Mesozoic vegetation covering about a hundred thousand square miles in the Rocky Mountains region. Europe and America now begin to show their modern contours.

It is important to notice that this great uprise of the land and the series of disturbances it entails differ from those which we summed up in the phrase Permian Revolution. The differences may help us to understand some of the changes in the living population. The chief difference is that the disturbances are more local, and not nearly simultaneous. There is a considerable emergence of land at the end of the Jurassic, then a fresh expansion of the sea, then a great rise of mountains at the end of the Cretaceous, and so on. We shall find our great mountain-masses (the Pyrenees, Alps, Himalaya, etc.) rising at intervals throughout the whole of the Tertiary Era. However, it suffices for the moment to observe that in the latter part of the Mesozoic and early part of the Tertiary there were considerable upheavals of the land in various regions, and that the Mesozoic Era closed with a very much larger proportion of dry land, and a much higher relief of the land, than there had been during the Jurassic period. The series of disturbances was, says Professor Chamberlin, "greater than any that had occurred since the close of the Palaeozoic."

From the previous effect of the Permian upheaval, and from the fact that the living population is now similarly annihilated or reduced, we should at once expect to find a fresh change in the climate of the earth. Here, however, our procedure is not so easy. In the Permian age we had solid proof in the shape of vast glaciated regions. It is claimed by continental geologists that certain early Tertiary beds in Bavaria actually prove a similar, but smaller, glaciation in Europe, but this is disputed. Other beds may yet be found, but we saw that there was not a general upheaval, as there had been in the Permian, and it is quite possible that there were few or no ice-fields. We do not, in fact, know the causes of the Permian icefields. We are thrown upon the plant and animal remains, and seem to be in some danger of inferring a cold climate from the organic remains, and then explaining the new types of organisms by the cold climate. This, of course, we shall not do. The difficulty is made greater by the extreme disinclination of many recent geologists, and some recent botanists who have too easily followed the geologists, to admit a plain climatic interpretation of the facts. Let us first see what the facts are.

In the latter part of the Jurassic we find three different zones of Ammonites: one in the latitude of the Mediterranean, one in the latitude of Central Europe, and one further north. Most geologists conclude that these differences indicate zones of climate (not hitherto indicated), but it cannot be proved, and we may leave the matter open. At the same time the warm-loving corals disappear from Europe, with occasional advances. It is said that they are driven out by the disturbance of the waters, and, although this would hardly explain why they did not spread again in the tranquil chalk-ocean, we may again leave the point open.

In the early part of the Cretaceous, however, the Angiosperms (flowering plants) suddenly break into the chronicle of the earth, and spread with great rapidity. They appear abruptly in the east of the North American continent, in the region of Virginia and Maryland. They are small in stature and primitive in structure. Some are of generalised forms that are now unknown; some have leaves approaching those of the oak, willow, elm, maple, and walnut; some may be definitely described as fig, sassafras, aralia, myrica, etc. Eastern America, it may be recalled, is much higher than western until the close of the Cretaceous period. The Angiosperms do not spread much westward; they appear next in Greenland, and, before the middle of the Cretaceous, in Portugal. They have travelled over the North Atlantic continent, or what remains of it. The process seems very rapid as we write it, but it must be remembered that the first half of the Cretaceous period means a million or a million and a half years.

The cycads, and even the conifers, shrink before the higher type of tree. The landscape, in Europe and America, begins to wear a modern aspect. Long before the end of the Cretaceous most of the modern genera of Angiosperm trees have developed. To the fig and sassafras are now added the birch, beech, oak, poplar, walnut, willow, ivy, mulberry, holly, laurel, myrtle, maple, oleander, magnolia, plane, bread-fruit, and sweet-gum. Most of the American trees of to-day are known. The sequoias (the giant Californian trees) still represent the conifers in great abundance, with the eucalyptus and other plants that are now found only much further south. The ginkgoes struggle on for a time. The cycads dwindle enormously. Of 700 specimens in one early Cretaceous deposit only 96 are Angiosperms; of 460 species in a later deposit about 400 are Angiosperms. They oust the cycads in Europe and America, as the cycads and conifers had ousted the Cryptogams. The change in the face of the earth would be remarkable. Instead of the groves of palm-like cycads, with their large and flower-like fructifications, above which the pines and firs and cypresses reared their sombre forms, there were now forests of delicate-leaved maples, beeches, and oaks, bearing nutritious fruit for the coming race of animals. Grasses also and palms begin in the Cretaceous; though the grasses would at first be coarse and isolated tufts. Even flowers, of the lily family (apparently), are still detected in the crushed and petrified remains.

We will give some consideration later to the evolution of the Angiosperms. For the moment it is chiefly important to notice a feature of them to which the botanist pays less attention. In his technical view the Angiosperm is distinguished by the structure of its reproductive apparatus, its flowers, and some recent botanists wonder whether the key to this expansion of the flowering plants may not be found in a development of the insect world and of its relation to vegetation. In point of fact, we have no geological indication of any great development of the insects until the Tertiary Era, when we shall find them deploying into a vast army and producing their highest types. In any case, such a view leaves wholly unexplained the feature of the Angiosperms which chiefly concerns us. This is that most of them shed the whole of their leaves periodically, as the winter approaches. No such trees had yet been known on the earth. All trees hitherto had been evergreen, and we need a specific and adequate explanation why the earth is now covered, in the northern region, with forests of trees which show naked boughs and branches during a part of the year.