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.
The majority of palaeontologists conclude at once, and quite confidently,
from this rise and spread of the deciduous trees, that a winter season
has at length set in on the earth, and that this new type of vegetation
appears in response to an appreciable lowering of the climate. The
facts, however, are somewhat complex, and we must proceed with caution.
It would seem that any general lowering of the temperature of the
earth ought to betray itself first in Greenland, but the flora of
Greenland remains far "warmer," so to say, than the flora
of Central Europe is to-day. Even toward the close of the Cretaceous
its plants are much the same as those of America or of Central Europe.
Its fossil remains of that time include forty species of ferns, as
well as cycads, ginkgoes, figs, bamboos, and magnolias. Sir A. Geikie
ventures to say that it must then have enjoyed a climate like that
of the Cape or of Australia to-day. Professor Chamberlin finds its
flora like that of "warm temperate" regions, and says that
plants which then flourished in latitude 72 degrees are not now found
above latitude 30 degrees.
There are, however, various reasons to believe that it is unsafe to
draw deductions from the climate of Greenland. There is, it is true,
some exaggeration in the statement that its climate was equivalent
to that of Central Europe. The palms which flourished in Central Europe
did not reach Greenland, and there are differences in the northern
Molluscs and Echinoderms which--like the absence of corals above the
north of England--point to a diversity of temperature. But we have
no right to expect that there would be the same difference in temperature
between Greenland and Central Europe as we find to-day. If the warm
current which is now diverted to Europe across the Atlantic--the Gulf
Stream--had then continued up the coast of America, and flowed along
the coast of the land that united America and Europe, the climatic
conditions would be very different from what they are. There is a
more substantial reason. We saw that during the Mesozoic the Arctic
continent was very largely submerged, and, while Europe and America
rise again at the end of the Cretaceous, we find no rise of the land
further north. A difference of elevation would, in such a world, make
a great difference in temperature and moisture.
Let us examine the animal record, however, before we come to any conclusion.
The chronicle of the later Cretaceous is a story of devastation. The
reduction of the cyeads is insignificant beside the reduction or annihilation
of the great animals of the Mesozoic world. The skeletons of the Deinosaurs
become fewer and fewer as we ascend the upper Cretaceous strata. In
the uppermost layer (Laramie) we find traces of a last curious expansion--the
group of horned reptiles, of the Triceratops type, which we described
as the last of the great reptiles. The Ichthyosaurs and Plesiosaurs
vanish from the waters. The "sea-serpents" (Mososaurs) pass
away without a survivor. The flying dragons, large and small, become
entirely extinct. Only crocodiles, lizards, turtle, and snakes cross
the threshold of the Tertiary Era. In one single region of America
(Puerco beds) some of the great reptiles seem to be making a last
stand against the advancing enemy in the dawn of the Tertiary Era,
but the exact date of the beds is disputed, and in any case their
fight is soon over. Something has slain the most formidable race that
the earth had yet known, in spite of its marvellous adaptation to
different environments in its innumerable branches.
We turn to the seas, and find an equal carnage among some of its most
advanced inhabitants. The great cuttlefish-like Belemnites and the
whole race of the Ammonites, large and small, are banished from the
earth. The fall of the Ammonites is particularly interesting, and
has inspired much more or less fantastic speculation. The shells begin
to assume such strange forms that observers speak occasionally of
the "convulsions" or "death-contortions" of the
expiring race. Some of the coiled shells take on a spiral form, like
that of a snail's shell. Some uncoil the shell, and seem to be returning
toward the primitive type. A rich eccentricity of frills and ornamentation
is found more or less throughout the whole race. But every device
--if we may so regard these changes--is useless, and the devastating
agency of the Cretaceous, whatever it was, removes the Ammonites and
Belemnites from the scene. The Mollusc world, like the world of plants
and of reptiles, approaches its modern aspect.
In the fish world, too, there is an effective selection in the course
of the Cretaceous. All the fishes of modern times, except the large
family of the sharks, rays, skates, and dog-fishes (Elasmobranchs),
the sturgeon and chimaera, the mud-fishes, and a very few other types,
are Teleosts, or bony-framed fishes--the others having cartilaginous
frames. None of the Teleosts had appeared until the end of the Jurassic.
They now, like the flowering plants on land, not only herald the new
age, but rapidly oust the other fishes, except the unconquerable shark.
They gradually approach the familiar types of Teleosts, so that we
may say that before the end of the Cretaceous the waters swarmed with
primitive and patriarchal cod, salmon, herring, perch, pike, bream,
eels, and other fishes. Some of them grew to an enormous size. The
Portheus, an American pike, seems to have been about eight feet long;
and the activity of an eight-foot pike may be left to the angler's
imagination. All, however, are, as evolution demands, of a generalised
and unfamiliar type: the material out of which our fishes will be
evolved.
Of the insects we have very little trace in the Cretaceous. We shall
find them developing with great richness in the following period,
but, imperfect as the record is, we may venture to say that they were
checked in the Cretaceous. There were good conditions for preserving
them, but few are preserved. And of the other groups of invertebrates
we need only say that they show a steady advance toward modern types.
The sea-lily fills the rocks no longer; the sea-urchin is very abundant.
The Molluscs gain on the more lowly organised Brachiopods.
To complete the picture we must add that higher types probably arose
in the later Cretaceous which do not appear in the records. This is
particularly true of the birds and mammals. We find them spreading
so early in the Tertiary that we must put back the beginning of the
expansion to the Cretaceous. As yet, however, the only mammal remains
we find are such jaws and teeth of primitive mammals as we have already
described. The birds we described (after the Archaeopteryx) also belong
to the Cretaceous, and they form another of the doomed races. Probably
the modern birds were already developing among the new vegetation
on the higher ground.
These are the facts of Cretaceous life, as far as the record has yielded
them, and it remains for us to understand them. Clearly there has
been a great selective process analogous to, if not equal to, the
winnowing process at the end of the Palaeozoic. As there has been
a similar, if less considerable, upheaval of the land, we are at once
tempted to think that the great selective agency was a lowering of
the temperature. When we further find that the most important change
in the animal world is the destruction of the cold-blooded reptiles,
which have no concern for the young, and the luxuriant spread of the
warm-blooded animals, which do care for their young, the idea is greatly
confirmed. When we add that the powerful Molluscs which are slain,
while the humbler Molluscs survive, are those which--to judge from
the nautilus and octopus--love warm seas, the impression is further
confirmed. And when we finally reflect that the most distinctive phenomenon
of the period is the rapid spread of deciduous trees, it would seem
that there is only one possible interpretation of the Cretaceous Revolution.
This interpretation--that cold was the selecting agency --is a familiar
idea in geological literature, but, as I said, there are recent writers
who profess reserve in regard to it, and it is proper to glance at,
or at least look for, the alternatives.
Before doing so let us be quite clear that here we have nothing to
do with theories of the origin of the earth. The Permian cold--which,
however, is universally admitted--is more or less entangled in that
controversy; the Cretaceous cold has no connection with it. Whatever
excess of carbon-dioxide there may have been in the early atmosphere
was cleared by the Coal-forests. We must set aside all these theories
in explaining the present facts.
It is also useful to note that the fact that there have been great
changes in the climate of the earth in past time is beyond dispute.
There is no denying the fact that the climate of the earth was warm
from the Arctic to the Antarctic in the Devonian and Carboniferous
periods: that it fell considerably in the Permian: that it again became
at least "warm temperate" (Chamberlin) from the Arctic to
the Antarctic in the Jurassic, and again in the Eocene: that some
millions of square miles of Europe and North America were covered
with ice and snow in the Pleistocene, so that the reindeer wandered
where palms had previously flourished and the vine flourishes to-day;
and that the pronounced zones of climate which we find today have
no counterpart in any earlier age. In view of these great and admitted
fluctuations of the earth's temperature one does not see any reason
for hesitating to admit a fall of temperature in the Cretaceous, if
the facts point to it.
On the other hand, the alternative suggestions are not very convincing.
We have noticed one of these suggestions in connection with the origin
of the Angiosperms. It hints that this may be related to developments
of the insect world. Most probably the development of the characteristic
flowers of the Angiosperms is connected with an increasing relation
to insects, but what we want to understand especially is the deciduous
character of their leaves. Many of the Angiosperms are evergreen,
so that it cannot be said that the one change entailed the other.
In fact, a careful study of the leaves preserved in the rocks seems
to show the deciduous Angiosperms gaining on the evergreens at the
end of the Cretaceous. The most natural, it not the only, interpretation
of this is that the temperature is falling. Deciduous trees shed their
leaves so as to check their transpiration when a season comes on in
which they cannot absorb the normal amount of moisture. This may occur
either at the on-coming of a hot, dry season or of a cold season (in
which the roots absorb less). Everything suggests that the deciduous
tree evolved to meet an increase of cold, not of heat.
Another suggestion is that animals and plants were not "climatically
differentiated "until the Cretaceous period; that is to say,
that they were adapted to all climates before that time, and then
began to be sensitive to differences of climate, and live in different
latitudes. But how and why they should suddenly become differentiated
in this way is so mysterious that one prefers to think that, as the
animal remains also suggest, there were no appreciable zones of climate
until the Cretaceous. The magnolia, for instance, flourished in Greenland
in the early Tertiary, and has to live very far south of it to-day.
It is much simpler to assume that Greenland changed--as a vast amount
of evidence indicates--than that the magnolia changed.
Finally, to explain the disappearance of the Mesozoic reptiles without
a fall in temperature, it is suggested that they were exterminated
by the advancing mammals. It is assumed that the spreading world of
the Angiospermous plants somewhere met the spread of the advancing
mammals, and opened out a rich new granary to them. This led to so
powerful a development of the mammals that they succeeded in overthrowing
the reptiles.
There are several serious difficulties in the way of this theory.
The first and most decisive is that the great reptiles have practically
disappeared before the mammals come on the scene. Only in one series
of beds (Puerco) in America, representing an early period of the Tertiary
Era, do we find any association of their remains; and even there it
is not clear that they were contemporary. Over the earth generally
the geological record shows the great reptiles dying from some invisible
scourge long before any mammal capable of doing them any harm appears;
even if we suppose that the mammal mainly attacked the eggs and the
young. We may very well believe that more powerful mammals than the
primitive Mesozoic specimens were already developed in some part of
the earth--say, Africa--and that the rise of the land gave them a
bridge across the Mediterranean to Europe. Probably this happened;
but the important point is that the reptiles were already almost extinct.
The difficulty is even greater when we reflect that it is precisely
the most powerful reptiles (Deinosaurs) and least accessible reptiles
(Pterosaurs, Ichthyosaurs, etc.) which disappear, while the smaller
land and water reptiles survive and retreat southward-- where the
mammals are just as numerous. That assuredly is not the effect of
an invasion of carnivores, even if we could overlook the absence of
such carnivores from the record until after the extinction of the
reptiles in most places.
I have entered somewhat fully into this point, partly because of its
great interest, but partly lest it be thought that I am merely reproducing
a tradition of geological literature without giving due attention
to the criticisms of recent writers. The plain and common interpretation
of the Cretaceous revolution--that a fall in temperature was its chief
devastating agency--is the only one that brings harmony into all the
facts. The one comprehensive enemy of that vast reptile population
was cold. It was fatal to the adult because he had a three-chambered
heart and no warm coat; it was fatal to the Mesozoic vegetation on
which, directly or indirectly, he fed; it was fatal to his eggs and
young because the mother did not brood over the one or care for the
other. It was fatal to the Pterosaurs, even if they
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