I have said that development is a "natural" process. We employ this word for the familiar and everyday occurrence or thing; it does not imply that everything is known about the object or phenomenon, because science knows that complete and final knowledge is impossible. We say that it is natural for rain to fall to the earth, and we speak of the law of gravitation according to which this takes place as a natural principle, but it may not have occurred to many to inquire what makes rain fall and why do masses of matter everywhere behave toward one another in the consistent manner described by the law in question. Sunshine is natural, but we do not know why light travels as it does from the sun to the earth, and this is another question which, like the inquiry into the ultimate cause of the familiar and natural phenomenon of gravitation, has not yet been answered. But it is still regarded as natural for the rain to fall and for the sun to shine. In the same way does science view development, denoting it natural because it is an ordinary everyday matter. And we are under no more obligation to postulate supernatural control for the changing forms in the life-history of a chick or a cat than we need to assume that gravitation and the radiation of light demand immediate supernatural direction. The embryology of no form is fully understood or described or explained, but no intelligent person would be willing to assert that because complete knowledge is lacking, it is unnatural for organic transformation to take place during growth. Whatever may be the ultimate origin and nature of the directing powers behind gravitation and development and other phenomena, we have no concern with such matters because they cannot be handled by scientific methods and one belief about them is on the same plane with any other. Our task is to deal with the everyday phenomena of life and the production of living species.
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It is not necessary to go far afield to find an animal which will introduce us to the general principles of embryology. In the present instance as in the case of comparative anatomy almost any form will disclose the meaning of development, for animate nature is uniform and consistent in its methods of operation throughout its wide range. We shall begin with the familiar frog which every one knows is a product of a tadpole; passing on to the chick we will learn more facts that will enable us to formulate the main principle of comparative embryology in definite terms; we will then be prepared to extend our survey so as to include somewhat less familiar facts and animals that are even more significant than the first illustrations.
If we should visit a woodland pond in early spring, we would find somewhere among the leaves and sticks in the water large masses of a clear jellylike consistency enclosing hundreds of little black spheres about an eighth of an inch in diameter. These are the egg masses and eggs of a common frog. Watching them day by day we see the small one-celled egg spheres divide into more and more numerous portions which are the daughter-cells, destined to form by their products the many varied tissues and organs of the developing larva and adult frog. After three or four days the egg changes from its globular form into an oval or elliptical mass, and from one end of this a small knob projects to become a flattened waving tail a few days later. On the sides of the larger anterior portion shallow grooves make their appearance and soon break through from the throat or pharynx to the exterior as gill-slits. Shortly afterwards the little embryo wriggles out of its encasing coat of jelly, develops a mouth, and begins its independent existence as a small tadpole, with eyes, nasal and auditory organs, and all other parts that are necessary for a free life. Thus the one-celled egg has transformed into something that it was not at first, and in doing this it has proved the possibility and the reality of organic reconstruction.
The tadpole breathes by means of its gills, and it is at first entirely devoid of the lungs which the adult frog possesses and uses. When we speak of the larval respiratory organs as gills we imply that they are like the organs of a fish which have the same name; they are truly like those of fishes, for the blood-vessels which go to them are essentially the same as in the lower types and they are supported by simple skeletal rods like the gill-bars of the fish. In a word, they are the same things.
The animal feeds and grows during the months of its first summer, and hibernates the following winter; with the warmth of spring it revives and proceeds further along the course of its development. Near the base of the tail two minute legs grow out from the hinder part of the body, and while these are enlarging two front legs make their appearance a little behind the gills. The tadpole now rises more frequently to the surface where it takes small mouthfuls of air. Meanwhile great changes are effected inside the body where the various systems of fishlike organs become remodeled into amphibian structures. A sac is formed from the wall of the esophagus, and this enlarges and divides to form the two simple lungs. The legs increase in size, the tail dwindles more and more, the gills close up, and soon the animal hops out on land as a complete young frog. From this time on it breathes by means of its lungs instead of gills, even though it returns to the water to escape its foes, to seek its prey, and to hibernate in the mud of the lake bed during the winter months.
All these changes are familiar and natural, but until science places them and similar facts in their proper relations their significance is lost to us. The tadpole is essentially a fish in its general structure and mode of life, even though its heritage is such that it can develop into a higher animal. When it does become a frog it proves beyond a doubt that there is no impassable barrier between fishes and amphibia. Our earlier comparison of the structures of these two classes of vertebrates led to the conclusion that the latter had evolved from antecedents like the former, and had thus followed them upon the earth; now that sequence seems to have some connection with the method by which a tadpole, obviously not a fish but nevertheless actually fishlike, changes into a frog, a member of a higher class of vertebrates. This method is employed by developing frogs apparently because it follows the ancestral order of events, and because, so to speak, the only way a frog knows how to become a frog is to develop from an egg first into a fishlike tadpole and then to alter itself as its ancestors did during their evolution in the past. We begin to see, then, that in addition to the impressive fact of development itself, the mode of organic transformation is far more conclusive evidence of evolution, because it reveals an order of events which parallels the order established by comparative anatomy as the evolutionary sequence.
However it is well to review some of the changes by which a chick comes into existence before attempting to comprehend fully the fundamental principle of development that the tadpole's history discloses to us. The egg of a common fowl is certainly not a chick. Within the calcareous shell are two delicate membranes that enclose the white or albumen; within this, swung by two thickened cords of the albumen, is the yellow yolk ball enclosed by a proper membrane of its own. In the earliest condition, even before the albumen and the shell are added and before the egg is laid, on one side of the yolk-mass there is a tiny protoplasmic spot which is at first a single cell and nothing more. The hen's egg is relatively enormous, but nevertheless, like that of the frog, it starts upon its course of development as a single unitary biological element—a cell. During the earliest subsequent hours the first cell divides again and again to form a small disk upon the surface of the yolk. Soon the cells along the middle line of this small sheet become rearranged to make an obvious streak or band, and about this line a simple tube is constructed which is destined to become the future brain and spinal cord. The whole disk continues to enlarge by further division of its constituent elements so that it encloses more and more of the yolk mass, but the little chick itself is made out of the cells along the central line of the original plate, from which it folds at the sides and in front and behind so as to lie somewhat above and apart from the flatter enclosing cell layers which partly surround the yolk.
At the sides of the primitive nerve-tube small blocks of cells arise to develop into primitive muscles and other structures. As nourishment is brought to the embryo from the surrounding layers enclosing the nutrient yolk, one system after another takes its shape and builds its several parts into organs which can be recognized as elementary structures of a chick. Among the more interesting ones are small clefts or slits formed in the side walls of the rudimentary throat or pharynx. Blood-vessels go forward from the simple heart to run up through the intervening bars exactly as in the tadpole and the fish. In brief, the young chick possesses a series of gill-slits, for these structures are the same in essential plan and relations as the clefts of tadpoles and fishes. Does this mean that even birds have descended from gill-breathing ancestors? Science answers in the affirmative, because evolution gives the only reasonable explanation of such facts as these. The case seems different from that of the frog, because gills are used by the tadpole, but gill-slits and gill-bars can have no conceivable value for the chick as organs concerned with the purification of the blood. None the less, if the transition from a gilled tadpole to the adult with lungs means an evolution of amphibia from fishlike ancestors, then the change of a chick embryo with gill-clefts into the fledgling without them is most reasonably interpreted as proof that birds as well as amphibia have had ancestors as simple as fishes.
As development progresses four small pads make their appearance; two of these lie on either side of the body back of the head and the other two arise near the posterior end. They are far from being wings and legs, but as day follows day they become molded into somewhat similar limbs, as much alike in general plan as the four legs of a lizard; subsequently the ones at the front change into real wings and the hinder ones become legs. Meanwhile the internal organs slowly transform from fishlike structures into things that display the characteristics of reptilian counterparts, and only later do they become truly avian. Last of all the finishing touches are made, and the whole creature becomes a particular kind of a bird which picks its way out of the shell and shifts for itself as a chick.
Only a few of the countless details have been mentioned which demonstrate the resemblance of the successive stages first to fishes, and later to amphibia and reptiles. We have a wide choice of materials, but even the foregoing brief list of illustrations shows that the order in which the stages follow is the one which comparative anatomy independently proves to be the order of the evolution of fishes, amphibia, reptiles, and birds. Why, now, should it be necessary for a developing bird to follow this order? The answer has been found in the immense array of embryological facts that investigators have verified and classified, that all tell the same story. It is, that birds have arisen by evolution from ancestors which were really as simple as the members of these lower classes. It seems then that the only way a bird of to-day can become itself is to traverse the path along which its progenitors had progressed in evolution. Stating its conclusions precisely, science formulates the principle in the following words: individual development is a brief resume of the history of the species in past times, or, more technically, ontogeny recapitulates phylogeny. To be sure, the full history is not reviewed in detail, for the chick embryo does not actually swim in water and breathe by means of gills. Only a condensed account of evolution of its kind is presented by an embryo during its development; as Huxley and Haeckel have put it, whole lines and paragraphs and even pages are left out; many false passages of a later date are inserted as the result of peculiar larval and embryonic needs and adjustments. But in its major statements and as a general outline, the account is a trustworthy natural document submitted as evidence that higher species of to-day have evolved from ancestors which must have been like some of the present lower animals.
Coming now to the mammalia, it might seem that we have reached forms so highly developed that they would not exhibit the same kind of developmental history, but would have their own mode of growing up. This is not so, for like the adult fish, the larval tadpole, and the embryo chick, an embryo of a cat or a man is at one time constructed with a series of gill-clefts and with blood-vessels and skeletal supports of fishlike nature that are everywhere associated with gills. The embryos of wildcats and dogs, rabbits and rats, pigs, deer, and sheep, and of all other mammalia, possess similar structures. Thus they all pass through a stage which is found also in the development of reptiles, birds, and amphibia,—a stage which corresponds to the fish throughout its life. Unless these facts mean that the great classes of vertebrates have originated together from the same or closely similar ancestors, they are unintelligible; for we cannot see why a cat or a chick should have to be essentially fishlike at any time unless this is so. Comparative anatomy states as we have learned that the amphibia as a class have evolved from and have out-developed the fishes, that reptiles have progressed still higher, and that birds and mammals have originated from reptilian ancestors along roads that have diverged beyond the immediate parent class. Because the members of each class have to pass along the same path trodden by their many varied ancestors, although at express speed, as it were, the similarity of the earliest stages in their development is explained, for during these periods they are traversing a path over which their ancestors passed together.
The places where the developing embryos depart from the common mode show where the several divisions took leave of one another in their evolution,—a point that comes out with great clearness when the facts of mammalian development are broadly compared. The embryos of carnivora and rodents and hoofed animals are alike in their earlier development, and their agreement means a community of origin. At a certain point the cat and dog depart from the common mode, but they remain alike up to a far later stage than the one in which they are similar to the embryos of rats and sheep. The rat and squirrel and rabbit, on their part, remain together until long after they take leave of the carnivora and ungulates; while the sheep and cattle and pigs have their own branch line, which they follow in company after leaving the embryos of the other orders. The reasons for these facts seem to be that the members of the three orders exemplified have evolved from the same stock, which accounts for their embryonic similarity for a long time after they collectively come to differ from amphibia and reptiles, while the members in each order became differentiated only later, wherefore their embryonic paths coincide for a longer period. Thus the degree of adult resemblance which indicates the closeness of relationship corresponds with the degree of embryonic agreement; that is, the cat and dog are much alike and their modes of development are essentially the same to the latest stages, while the cat and horse agree only during the earliest and middle stages, and their lines diverge before those of the cat and dog on the one hand, or those of the horse and pig on the other.
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Like the fundamental principle of comparative anatomy in its sphere, the Law of Recapitulation, formulated as a summary description of the foregoing and similar facts, is one that holds true throughout the entire range of embryology and for every division of the animal series, however large or small. We have discussed its broader application, and now we may take up some of the more or less special cases mentioned in the earlier section of the present chapter, to see how it may work in detail.
The flounder was noted as a variant of the fish theme which seemed to be a descendant of a symmetrical ancestor because its structural plan was like that of other bony fishes. If this be true, and if in its development a flounder must review its mode of evolution as a species, the young fish ought to be symmetrical; and it actually is. The grotesque skate and hammerhead shark were demonstrated to be derivatives of a simpler type of shark; their embryos are practically indistinguishable from those of ordinary dogfish and sharks.
Among the jointed animals a wealth of interesting material is found by the embryologist. All crabs seemed to be modified lobsterlike creatures; to confirm this interpretation, based solely upon details of adult structure, young crabs pass through a stage when to all intents and purposes they are counterparts of lobsters. Even the twisted hermit crab, which has a soft-skinned hinder part coiled to fit the curve of the snail shell used as a protection, is symmetrical and lobster-like when it is a larva.
Among the insects many examples occur that are already familiar to every one. The egg of a common house-fly hatches into a larva called a maggot; in this condition the body destined to become the vastly different fly is composed of soft-skinned segments very much alike and also similar to the joints of a worm. Comparative anatomy demonstrates that the fly and all other insects have arisen from wormlike ancestors, whose originally similar segments later differentiated in various ways to become the diverse segments of adult insects; the embryonic history of flies of to-day corroborates these assertions, in so far as every individual fly actually does become a wormlike larva before it changes into the final and complete adult insect. The other kinds of insects are equally striking in their life-histories. All beetles, such as the potato bug and June bug, develop from grubs which, like the maggots of flies, are similar to worms in numerous respects. Butterflies and moths pass through a caterpillar stage having even more striking resemblances to worms. All the larvae of insects are therefore like one another, and like worms also, in certain fundamental characters of internal and external structure; so the conclusion that the whole group of insects has arisen by evolution from more primitive ancestors resembling the worms of to-day is based upon mutually explanatory details of comparative anatomy and embryology.
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Let us now turn back to some of the earlier pages of the embryological record which we passed over in order that we might translate the later portions dealing with more familiar and intelligible structures like gills. Before the egg of the frog becomes an elliptical mass of cells, it is at one time a double-walled sac enclosing a central cavity; in this stage it is called a gastrula. Tracing back the mode of its formation, we find that it is produced from a hollow sphere of fewer cells that are essentially alike; this stage also is so important that the special term blastula is applied to it. Still earlier, there are fewer cells—128 or thereabouts, 64, 32, 16, 8, 4, 2, and 1. In other words, the starting point in the development of the frog is a single biological unit; this divides and its products redivide to constitute the many-celled blastula and the double-walled gastrula. All the other animals we have mentioned begin like the frog, as eggs which are single cells and nothing more; they too pass on to become blastulae and gastrulae, similar to those of the frog in all essential respects, particularly as regards the nature of the organs produced by each of the two primary layers, and the mode of their formation. Does the occurrence of blastulae and gastrulae and one-celled beginnings mean that the higher animals composed of numerous and much differentiated cells have evolved in company from two-layered saccular ancestors which were themselves the descendants of spherical colonies of like cells, and ultimately of one-celled animals?
Comparative anatomy has asserted that this is so, as we have already learned, for it finds that adult animals array themselves at different levels of a scale beginning at the bottom with the protozoa, continuing on to the two-layered animals like Hydra and jellyfish and sea-anemones, and then extending upwards to the region of the more complicated invertebrates and vertebrates. It was difficult perhaps to believe that these successive grades of organic structure indicated an order of evolution, because it seemed impossible that an animal so simple as a protozoan could produce offspring with the complex organization of a frog or a cat, even in long ages. But development delivers its evidence relating to this matter with telling and impressive force. How can we doubt the possibility of an evolution of higher animals from ancestors as simple as Hydra and Amoeba when a frog and a cat, like all other complicated organisms, begin individual existence as single cells, and pass through gastrula stages? If we deny it, we contradict the evidence of our senses, for the development is actually accomplished by the transformation of a single cell into a double-walled sac, and of this into different and more intricate organic mechanisms. The process can take place, for it does take place. Not until the investigator becomes familiar with a wide range of diverse animals and the peculiar qualities of their similar early stages, can he estimate the tremendous weight of the facts of comparative embryology. Were the statement iterated and reiterated on every page and in every paragraph, there would be no undue emphasis put upon the astounding fact that the apparently impassable gap between a one-celled animal like Amoeba and a mammal like a cat is actually compassed during the development of the last-named organisms from single cells. The occurrence of gill-slits in the embryos of lizards, birds, and mammals now seems a small thing when compared with the correspondences disclosed by the earliest stages of development. But in spite of their complexity, all the changes of "growing up" are explained and understood by the simple formula that the mode of individual development owes its nature primarily to the hereditary influence of earlier ancestors back to the original animals which were protozoa.
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Embryology as a distinct division of zooelogy has grown out of studies of classification and comparative anatomy. Its beginnings may be found in medieval natural history, for as far back as 1651 Harvey had pointed out that all living things originate from somewhat similar germs, the terse dictum being "Ex ovo omnia." By the end of the eighteenth century many had turned to the study of developing organisms, though their views by no means agreed as to the way an adult was related to the egg. Some, like Bonnet, held that the germ was a minute and complete replica of its parent, which simply unfolded and enlarged like a bud to produce a similar organism. Even if this were true, little would be gained, for it would still remain unknown how the germinal miniature originated to be just what it was conceived and assumed to be. Wolff was the originator of the view that is now practically universal among naturalists, namely, that development is a real process of transformation from simpler to more complex conditions.
The subject of comparative embryology grew rapidly during the nineteenth century as the field of comparative anatomy became better known, and when naturalists became interested in animals, not only as specific types, but also as the finished products of an intricate series of transformations. When life-histories were more closely compared, the meaning of the resemblances between early stages of diverse adult organisms was read by the same method which in comparative anatomy finds that consanguinity is expressed by resemblance. The great law of recapitulation, stated in one form by Von Baer and more definitely by Haeckel in the terms employed in the foregoing sections, was for a time too freely used and too rigidly applied by naturalists whose enthusiasm clouded their judgment. A strong reaction set in during the latter part of the nineteenth century, when attention was directed to the anachronisms of the embryonic record and to the alterations that are the results of larval or embryonic adaptation as short cuts in development. Nevertheless, it is not seriously questioned, I believe, that the main facts of a single life-history owe their nature to the past evolution of the species to which a given animal belongs.
Nowadays the problems in this well-organized department are concerned not only with more accurate accounts of the development of animals, but also with the mechanics of development, with the relative value of external and internal influences, and above all with the physical basis of inheritance. It is clear that the factors that direct the development of a wood frog's egg so that it becomes a wood-frog and not a tree-toad must lie in the egg itself, as derivatives from the two parent organisms. Weismann and his followers have proved that a peculiar substance in the nuclei of the egg and its daughter-products contains the essential factors of development, whatever these may be. Experiments dealing with the phenomena of heredity in pure and mixed breeds have largely confirmed Weismann's doctrine, and they have prepared the way for a deeper investigation of the marvelous process of biological inheritance.
However much he may be interested in the details of embryological science, the general student of natural history is more concerned with the bearing of its primary laws upon the great problem of evolution. In the foregoing brief review of the fundamental facts and principles of this subject, the purpose has been to show how the phenomena of development are viewed by men of science, and how they take their place in the doctrine of organic evolution. And it has also been made plain that comparative anatomy and comparative embryology support and supplement one another in countless ways and places, although each in itself is a complete demonstration that evolution is a real and a natural process.
THE EVIDENCE OF FOSSIL REMAINS
Few natural objects appeal to the interest and imagination of the student with more force than the fragments of animals and plants released from the rocks where they have been entombed for ages. Our lives are so brief that it is impossible for us to comprehend the full duration of the slow process which constructed the burial shrouds of these creatures of long ago. We try to picture the earth and its inhabitants as they were when lizards were the highest forms of animals, and we wonder how life was lived in the dense forests of the coal age. Science can never learn all about the ancient history of the earth and of the organisms of bygone times; yet it has been able to accomplish much through its endeavors to reconstruct the past, for its method is one by which sure results can always be obtained whenever there are definite facts with which it can work. In our present study of evolution we reach the point when we must examine the testimony of the rocks, and the results and methods of that department of knowledge called palaeontology, which is concerned with fossils and their interpretation.
The word "palaeontology" means literally the "science of living things of long ago." It deals directly with the remains of animals and plants found as fossils, and it interprets them through its knowledge of the way modern animals are constructed and of the changes the earth's crust has undergone. A skull-like object may be found in a coal field and may come into the hands of the palaeontologist: from his acquaintance with the head skeletons of recent types he will be able to assign the extinct creature which possessed the skull to a definite place in the animal scale and to understand its nearer or wider affinities with other animals of later times and of earlier epochs. In doing these things palaeontology employs the methods of comparative anatomy with which we have now become familiar. In the performance of its other tasks, however, palaeontology must work independently. It is necessary to know when a fossilized animal lived, not that its time need be measured by an absolute number of a few thousands or millions of years antedating our own era, for that is impossible. But the important thing is to know its relative age, and whether it preceded or followed other similar animals of its own group or of different divisions. The rocks themselves must be understood, how they have been formed and how they are related in mineralogical nature and in historical succession. Palaeontology also deals with a number of subjects that are not in themselves biological, such as the combination of circumstances necessary for the adequate preservation of fossil relics. In so far as it is concerned with physical matters, as contrasted with strictly biological data, it is one with geology. Indeed, the investigators in these two departments must always work side by side and render mutual assistance to one another in countless ways, for each division needs the results of the other in order to accomplish its own distinct purposes. It must be evident to every one that it is impossible to understand the meaning of fossils and the place of the testimony of the rocks in the doctrine of evolution without knowing much about the geological history of the earth and the influences at work in the past. For these reasons palaeontology differs somewhat from the other divisions of zooelogy where direct observation gives the materials for arrangement and study; in this case the individual data, that is, the fossil fragments themselves, can be made available only through a knowledge of their exact situations, of the reasons for their occurrence in particular places in the rock series and of the way rocks themselves are constructed and worked over by natural agencies. Our task is therefore twofold: certain physical matters of a geological nature must first be investigated before the biological facts can be described.
No doubt most people feel justified in believing that the whole doctrine of evolution must stand or fall according to the cogency of the palaeontological evidences. Plain common sense says that the owners of shelly or bony fragments found in the deeply-laid strata of the earth must have lived countless years ago, and if the evolutionist asserts that primitive organic forms of ancient times have produced changed descendants of later times, it would seem that fossil evidence would be supremely and overwhelmingly important. It is true, of course, that this evidence is peculiarly significant, because in some ways it is more direct than that of the other categories already outlined. But it must not be forgotten that the doctrine is already securely founded upon the basic principles of anatomy and embryology. Science must treat the data of this category by different methods and must view them in different ways. Therefore we are interested in palaeontology because of the way it tells the story of evolution in its own words, and because we are justified in expecting that its account should include a description of some such order of events as that revealed by the developing embryos of modern organisms and that demonstrated by the comparative anatomy of the varied species of adult animals.
It is true that palaeontology gives direct testimony about the evolutionary succession of animals in geologic time. But we now know that embryology is even more direct in its proof that organic transformation is natural and real; while at the same time there is a completeness in the full series of developmental stages connecting the one-celled egg with the adult creature that must be forever lacking in the case of the fossil sequence of species. If paragraphs and pages are missing from the brief embryonic recapitulation, whole chapters and volumes of the fossil series have been lost for all time. The investigators whose task it has been to decipher the story of the earth's evolution have had to meet numerous and exasperating difficulties which do not confront the embryologist and anatomist who study living materials. Nevertheless the library of palaeontological documents is one which has been founded for over a century, and it has grown fast during recent decades, so that consistent accounts may now be read of the great changes in organic life as the earth has altered and grown older. And in all this record, there is not a single line or word of fact that contradicts evolution. What definite evidence there is tells uniformly in favor of the doctrine, for it is possible, in the first place, to work out the order of succession of many of the great groups of animals, and this order is found to be the same as that established by the other bodies of evidence. Secondly, some fossil groups are astonishingly complete, so that the ancient history of a form like the horse can be written with something approaching fullness. Finally, the remains of certain animals have been found so situated in geological ways, and so constructed anatomically, that the zooelogist is justified in denoting them "missing links," because they seem to have been intermediate between groups that have diverged so widely during recent epochs as to render their common ancestry scarcely credible.
With these general results in mind, we must now become acquainted with such subjects as the interpretation of fossils, the causes for the incompleteness of the series, the conditions for fossilization, the forces of geological nature, and other matters that make the fossils themselves intelligible as scientific evidence.
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Many views have been entertained regarding the actual nature of the relics of antiquity exhumed from the rocks or exposed upon the surface by the wear and tear of natural agencies. In earliest times such things were variously considered as curious freaks of geological formation, as sports of nature, or as the remains of the slain left upon the battle-ground of mythical Titans. Some of the Greeks supposed that fossils were parts of animals formed in the bowels of the earth by a process of spontaneous generation, which had died before they could make their way to the surface. They were sometimes described as the bones of creatures stranded upon the dry land by tidal waves, or by some such catastrophe as the traditional flood of the scriptures. In medieval times, and even in our own day, some people who have been opposed to the acceptance of any portion of the doctrine of evolution have actually defended the view that the things called fossils were never the shells or bones of animals living in bygone times, but that they only simulate such things and have been created as such together with the layers of rock from which they may have been taken. If we employed the same arguments in dealing with the broken fragments of vases and jewelry taken from the Egyptian tombs or from the buried ruins of Pompeii, we would have to believe that such pieces were created as fragments and that they were never portions of complete objects, just because no one alive to-day has ever seen the perfect vessel or bracelet fashioned so long ago. Common sense directs us to discard such a fantastic interpretation in favor of the view that fossils are what they seem to be—simply relics of creatures that lived when the earth was younger.
Until this common sense view was adopted there was no science of palaeontology. Cuvier was the first great naturalist to devote particular attention to the mainly unrelated and unverified facts that had been discovered before his time. He was truly the originator of this branch of zooelogy, for he brought together the observations of earlier men and extended his own studies widely and surely, emphasizing particularly the necessity for noting carefully the geological situation of a fossil in rocks of an older or later period of formation. His great result was the demonstration that many groups of animals existed in earlier ages that seem to have no descendants of the same nature to-day, and also that many or most of our modern groups are not represented in the earliest formed sedimentary rocks, although these recent forms possess hard parts which would surely be present somewhere in these levels if the animals actually existed in those times. But the meaning of these facts escaped Cuvier's mind. He was a believer in special creation, like Linnaeus and all but a few among his predecessors, and he explained the diversity of the faunas of different geological times in what seems to us a very simple and naive way. In the beginning, he held, when the world was created, it was furnished with a complete set of animals and plants. Then some great upheaval of nature occurred which overwhelmed and destroyed all living creatures. The Creator then, in Cuvier's view, proceeded to construct a new series of animals and plants, which were not identical with those of the former time, but were created according to the same general working plans or architectural schemes employed before. Another cataclysm was supposed to have occurred, which destroyed the second series of organisms and laid a new covering of rocks over the earth's surface for a subsequent period of relative quiet; and so the process was continued. By this account, Cuvier endeavored to reconcile the doctrine of supernatural creation and intervention with the obvious facts that organisms have differed at various times in the earth's history. Although he saw that animals of successive periods displayed similar structures, like the skeleton of vertebrates, which testified to some connection, Cuvier could not bring himself to believe that this connection was a genealogical one.
Mainly through the influence of the renowned English man of science, Charles Lyell, the students of the earth came to the conclusion that its manifold structures had developed by a slow and orderly process that was entirely natural; for they found no evidence of any sudden and drastic world-wide remodeling such as that postulated by the Cuvierian hypothesis of catastrophe. The battle waged for many years; but now naturalists believe that the forces, of nature, whose workings may be seen on all sides at the present time, have reconstructed the continents and ocean beds in the past in the same way that they work to-day. The long name of "uniformitarianism" is given to Lyell's doctrine, which has exerted an influence upon knowledge far outside the department of geology. Darwin tells us how much he himself was impressed by it, and how it led him to study the factors at work upon organic things to see if he could discern evidence of a biological uniformitarianism, according to which the past history of living things might be interpreted through an understanding of their present lives.
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What, now, are the reasons why the palaeontological evidence is not complete and why it cannot be? In the first place the seeker after fossil remains finds about three fifths of the earth's surface under water so that he cannot explore vast areas of the present ocean beds which were formerly dry land and the homes of now extinct animals. Thus the field of investigation is seriously restricted at the outset, but the naturalist finds his work still more limited, in so far as much of the dry land itself is not accessible. The perennial snows of the Arctic region render it impossible to make a thorough search in the frigid zone, and there are many portions of the temperate and torrid zones that are equally unapproachable for other reasons. But even where exploration is possible, the surface rocks are the only ones from which remains can be readily obtained, for the layers formed in earlier ages are buried so deeply that their contents must remain forever unknown in their entirety. Only a few scratches upon the earth's hard crust have been made here and there, so it is small wonder that the complete series of extinct organisms has not been produced by the palaeontologist.
A brief survey of the varied groups of animals themselves is sufficient to bring to light many biological reasons which account for still more of the vacant spaces in the palaeontological record. We would hardly expect to find remains of ancient microscopic animals like the protozoa, unless they possessed shells or other skeletal structures which in their aggregate might form masses like the chalk beds of Europe. Jellyfish and worms and naked mollusks are examples of the numerous orders of lower animals having no hard parts to be preserved, and so all or nearly all of the extinct species belonging to these groups can never be known. But when an animal like a clam dies its shell can resist the disintegrating effects of bacteria and other organic and inorganic agencies which destroy the soft parts, and when a form like a lobster or a crab, possessing a body protected by closely joined shell segments, falls to the bottom of the sea, the chances are that much of the animal's skeleton will be preserved. Thus it is that corals, crustacea, insects, mollusks, and a few other kinds of lower forms constitute the greater mass of invertebrate palaeontological materials because of their supporting structures of one kind or another. Perhaps the skeletal remains of the vertebrates of the past provide the student of fossils with his best facts, on account of the resistant nature of the bones themselves, and because the backboned animals are relatively modern; then, too, the rocks in which their remains occur have not been so much altered by geological agencies, or buried so deeply under the strata formed later. Of course only the hardest kinds of shells would remain as such after their burial in materials destined to turn into rock; in the majority of cases, an entombed bone is infiltrated or replaced by various mineral substances so that in time little or nothing of the original thing would remain, though a mold or a cast would persist.
But even if an animal of the past possessed hard structures, it must have satisfied certain limited conditions to have its remains prove serviceable to students of to-day. A dead mammal must fall upon ground that has just the right consistency to receive it; if the soil is too soft, its several parts will be separated and scattered as readily as though it had fallen upon hard ground where it would be torn to pieces by carnivorous animals. The dead body must then be covered up by a blanket of silt or sand like that which would be deposited as the result of a freshet. If a skeleton is too greatly broken up or scattered, it may be difficult or even impossible for its discoverer to piece together the various fragments and assemble them in their original relations. Very few individuals have been so buried and preserved as to meet the conditions for the formation of an ideal fossil. To realize how little may be left of even the most abundant of higher organisms, we have only to recall that less than a century ago immense herds of bison and wild horses roamed the Western plains, but very few of their skulls or other bones remain to be enclosed and fossilized in future strata of rocks. When we appreciate all these difficulties, both geological and biological, we begin to see clearly why the ancient lines of descent cannot be known as we know the path and mode of embryonic transformation. The wonder is not that the palaeontological record is incomplete, but that there is any coherent and decipherable record at all. Yet in view of the many and varied obstacles that must be surmounted by the investigator, and the adverse factors which reduce the available evidence, the rapidly growing body of palaeontological facts is amply sufficient for the naturalist to use in formulating definite and conclusive principles of evolution.
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For the purposes of palaeontology, the most essential data of geology are those which indicate the relative ages of the strata that make up the hard outer crust of the earth, for only through them can the order of animal succession be ascertained. It does not matter exactly how old the earth may be. While it is possible to determine the approximate length of time required for the construction of sedimentary rocks like those which natural agencies are producing to-day, there are few definite facts to guide speculation as to the mode or duration of the process by which the first hard crystalline surface of the earth was formed. But palaeontology does not care so much about the earliest geological happenings, for it is concerned with the manifold animal forms that arose and evolved after life appeared on the globe. Questions as to the way life arose, and as to the earliest transformations of the materials by which the earth was first formed are not within the scope of organic evolution, although they relate to intensely interesting problems for the student of the process of cosmic evolution.
According to the account now generally accepted, the original material of the earth seems to have been a semi-solid or semi-fluid mass formed by the condensation of the still more fluid or even gaseous nebula out of which all the planets of the solar system have been formed and of which the sun is the still fiery core. As soon as the earth had cooled sufficiently its substances crystallized and wrinkled to form the first mountains and ridges; between and among these were the basins which soon filled with the condensing waters to become the earliest lakes and oceans. The wear and tear of rains and snows and winds so worked upon the surfaces of the higher regions that sediments of a finer or coarser character like sand and mud and gravel were washed down into the lower levels. These sediments were afterwards converted into the first rocks of the so-called stratified or sedimentary series, as contrasted with the crystalline or plutonic rocks like the original mass of the earth and the kinds forced to the surface by volcanic eruptions. Later the earth wrinkled again in various ways and places so that new ridges and mountains were formed with new systems of lakes and oceans and rivers; and again the elements continued to erode and partially destroy the higher masses and to lay down new and later series of sedimentary rocks upon the old.
It seems scarcely credible that the apparently weak forces of nature like those we have mentioned are sufficiently powerful to work over the massive crust of the earth as geology says they have. Our attention is caught, as a rule, only by the greater things, like the earthquakes at San Francisco and Valparaiso, and the tidal waves and cyclones of the South Seas; but the results of these sporadic and local cataclysms are far less than the effects of the persistent everyday forces of erosion, each one of which seems so small and futile. When we look at the Rocky Mountains with their high and rugged peaks, it seems almost impossible that rain and frost and snow could ever break them up and wear them down so that they would become like the rounded hills of the Appalachian Mountain chain, yet this is what will happen unless nature's ways suddenly change to something which they are not now. A visitor to the Grand Canon of the Colorado sees a magnificent chasm over a mile in depth and two hundred miles long which has actually been carved through layer after layer of solid rock by the rushing torrents of the river. Perhaps it is easier to estimate the geological effects of a river in such a case as Niagara. Here we find a deep gorge below the famous falls, which runs for twenty miles or so to open out into Lake Ontario. The water passing over the brim of the falls wears away the edge at a rate which varies somewhat according to the harder or softer consistency of the rocks, but which, since 1843, has averaged about 104 inches a year. Knowing this rate, the length of the gorge, and the character of the rocky walls already carved out, the length of time necessary for its production can be safely estimated. It is about 30,000 to 40,000 years, not a long period when the whole history of the earth is taken into account. A similar length of time is indicated for the recession of the Falls of St. Anthony, of the Mississippi River, an agreement that is of much interest, for it proves that the two rivers began to make their respective cuttings when the great ice-sheet receded to the north at the end of the Glacial epoch.
What has become of the masses washed away during the formation of these gorges? As gravel and mud and silt the detritus has been carried to the still waters of the lower levels, to be laid down and later solidified into sandstone and slate and shale. All over the continents these things are going on, and indefatigable forces are at work that slowly but surely shear from the surface almost immeasurable quantities of earth and rock to be transported far away. In some instances it is possible to find out just how much effect is produced in a given period of time, especially in the case of the great river systems. For example, the mass of the fine particles of mud and silt carried in a given quantity of the water of the Mississippi as it passes New Orleans can be accurately measured, and a satisfactory determination can also be made of the total amount of water carried by in a year. From these figures the amount of materials in suspension discharged into the Gulf of Mexico becomes known. It is sufficient to cover one square mile to the depth of 269 feet; in twenty years it is one cubic mile, or five cubic miles in a century. Turning now to the other aspect of this process, and the antecedent causes which produce these effects, it appears that the area of the Mississippi River basin is 1,147,000 square miles—about one third of the total area of the United States. Knowing this, and the annual waste from its surface, it is easy to demonstrate that it will take 6000 years to plane off an average of one foot of soil and rock from the whole of this immense area. Of course only an inch or a few inches will be taken from some regions where the ground is harder or rockier, or where little rain falls, while many feet will be washed away from other places. The waters of the Hoang-ho come from about 700,000 square miles of country, from which one foot of soil is washed away in 1464 years. The Ganges River, draining about 143,000 square miles, carries off a similar depth of eroded materials from its basin in 823 years! Should we add to the above figures those that specify the bulk of the chemical substances in solution carried by these waters, the total would be even greater. We know that in the case of the Thames River, calcareous substances to the amount of 10,000 tons a year are carried past London, and all this mineral has been dissolved by rain-water from the chalky cliffs and uplands of England, so that the land has become less by this amount. Thus we learn that vast alterations are being made in the structure of great continents by rain and rivers, as well as by glaciers and other geological agencies. And at the same time that old strata are undergoing destruction new ones are in process of construction at other places, where animal remains can be embedded and preserved as fossils. The forces at work seem weak, but they continue their operations through ages that are beyond our comprehension and they accomplish results of world-building magnitude.
Thus the whole process of geological construction is such that older exposed strata continually undergo disintegration, but this involves the destruction of any fossils that they might contain. The very forces that preserve the relics of extinct animals at one time undo their work at a later period. There are many other influences besides that destroy the regularity of rock layers or change their mineralogical characters by metamorphosis. It is easier to see how volcanic outbursts alter their neighboring territory. The intense subterranean heat and imprisoned steam melt the deeper substances of the earth's crust, so that these materials boil out, as it were, where the pressure is greatest, and where lines of fracture and lesser resistance can be found. Because so much detritus is annually added to the ocean floors—enough to raise the levels of the oceans by inches in a century—it is natural that greater pressures should be exerted in these areas than in the slowly thinning continental regions. These are some of the reasons why volcanoes arise almost invariably along the shores or from the floors of great ocean beds. The chain that extends from Alaska to Chili within the eastern shore of the Pacific Ocean, and the many hundreds of volcanoes of the Pacific Islands bring to the surface vast quantities of eruptive rocks which break up and overlie the sedimentary strata formed regularly in other ways and at other times. The volcanoes of the Java region alone have thrown out at least 100 cubic miles of lava, cinders, and ashes during the last 100 years—twenty times the bulk of the materials discharged into the Gulf of Mexico by the Mississippi River in the same period of time.
From these and similar facts, the naturalist finds how agencies of the present construct new rocks and alter the old; and so in the light of this knowledge, he proceeds with his task of analyzing the remote past, confident that the same natural forces have done the work of constructing the lower geological levels because these earlier products are similar to those being formed to-day. After learning this much, he must immediately undertake to arrange the strata according to their ages. This might seem a difficult or even an impossible task, but the rocks themselves provide him with sure guidance.
Wherever a river has graven its deep way through an area of hard rocks, as in the case of Niagara, the walls display on their cut surfaces a series of lines and planes showing that they are superimposed layers formed serially by deposits that have differed some or much at different times according to the circumstances controlling the erosion of their constituent particles. A layer of several feet in thickness may be composed of compact shale, while above it will be a zone of limestone, and again above this another layer of shale. Successive strata like these, where they are parallel and obviously undisturbed, are evidently arranged in the order of their formation and age. But by far the most impressive demonstration of the basic principle of geology employed for the determination of the relative ages of rocks is the mighty Canon of the Colorado. As the traveler stands on the winding rim of this vast chasm, his eye ranges across 13 miles of space to the opposite walls, which stretch for scores of miles to the right and left; upon this serried face he will see zone after zone of yellow and red and gray rock arranged with mathematical precision and level in the same order as on the steep slopes beneath him. Plain common sense tells him that the great sheets of rock stretched continuously at one time between the now separate walls, and that the various strata of sandstone and limestone were deposited in successive ages from below upwards in the order of their exposure. When now he extends his explorations to another state like Utah or Wyoming, he may find some but not all of the series exhibited in the Grand Canon, overlaid or underlaid by other strata which in their turn can be assigned to definite places in the sequence. By the same method, the geologist correlates and arranges the rocks not only of different parts of the same state, or of neighboring states, but even those of widely separated parts of North America and of different continents. But he learns that he must refrain from over-hasty conclusions, for he soon finds that the sedimentary rocks have not been constructed at the same rate in different places during one and the same epoch, and that rocks formed even at one period are not always identical in nature. But his guiding principle is sensible and reasonable, and by employing it with due caution he provides the palaeontologist with the requisite knowledge for his special task, which is to arrange the extinct animals whose remains are found as fossils of various earth ages in the order of their succession in time.
CONDENSED TABLE OF PALAEONTOLOGICAL FACTS
YEARS NUMBER OF ORDER OF NECESSARY FOR FEET IN GEOLOGICAL GEOLOGICAL APPEARANCE OF FORMATION THICKNESS AGE EPOCH CHARACTERISTIC GROUPS M B R A F I a i e m i n b m r p p s v r Recent m d t h h e a or a s i i e r t Quaternary l l b s t e s e i e s s a - Pleistocene Cenozoic Pliocene 5,000,000 25,000 or Miocene Tertiary Oligocene Eocene Mesozoic Cretaceous 4,000,000 23,000 or Jurassic Secondary Triassic Permian Palaeozoic Carboniferous 21,000,000 106,000 or Devonian Primary Silurian Cambrian 20,000,000 30,000 Azoic Archaen
After what seems an unduly long preparation, we now come to the actual biological evidence of evolution provided by the results of this division of zooelogical science. But all of the foregoing is fundamentally part of this department of knowledge and it is absolutely essential for any one who desires to understand what the fossils themselves demonstrate.
The oldest sedimentary rocks are devoid of fossil remains and so they are called the Azoic or Archaean. They comprise about 30,000 feet of strata which seem to have required at least 20,000,000 years for their formation. This period is roughly two-fifths of the whole time necessary for the formation of all the sedimentary rocks, and this proportion holds true even if the entire period of years should be taken as 100,000,000 instead of 50,000,000 or less. The earth during this early age was slowly organizing in chemical and physical respects so that living matter could be and indeed was formed out of antecedent substances—but this process does not concern us here. The important fact is that the second major period, called the Palaeozoic, or "age of ancient animals," saw the evolution of the lowest members of the series,—the invertebrates,—and the most primitive of the backboned animals, like fishes and amphibia. The rocks of this long age include about 106,000 feet of strata, demanding some 21,000,000 or 22,000,000 years for their deposition. Thus it is proved that the invertebrate animals were succeeded in time by the higher vertebrates, which is exactly what the evidences of the previous categories have shown. When we remember that the lower animals are devoid as a rule of skeletal structures that might be fossilized, and when we recall the fact that the strata of the palaeozoic provided the materials out of which the upper layers were formed afterwards, we can understand why the ancient members of the invertebrate groups are not known as well as the later and higher forms like vertebrates. Yet all the fossils of these relatively unfamiliar creatures clearly prove that no complex animal appears upon a geological horizon until after some simple type belonging to a class from which it may have taken its origin; in brief, there are no anachronisms in the record, which always corresponds with the record written by comparative anatomy, wherever the facts enable a comparison to be made.
But the extinct animals of the third and fourth ages are more interesting to us, because there are more of them and because they are more like the well-known organisms of our present era. These two ages are called the Mesozoic or Secondary, and the Cenozoic or Tertiary. The former is so named because it was a transitional age of animals that are intermediate in a general way between the primitive forms of the preceding age and those of the next period; the latter name means the "recent-animal" age, when evolution produced not only the larger groups of our present animal series, but also many of the smaller branches of the genealogical tree like orders and families to which the species of to-day belong.
Confining our attention to the large vertebrate classes, the testimony of the rocks proves, as we have said, that fishes appeared first in what are called the Silurian and Devonian epochs, where they developed into a rich and varied array of types unequaled in modern times. At that period, they were the highest existing animals—the "lords of creation," as it were. To change the figure, their branch constituted the top of the animal tree of the time, but as other branches grew upwards to bear their twigs and leaves, as the counterparts of species, the species of the branch of fishes decreased in number and variety, as do the leaves of a lower part of a tree when higher limbs grow to overshadow them.
Following the fishes, the amphibia arose during the coal age or Carboniferous, usurping the proud position of the lower vertebrate class. The reptiles then appeared and gained ascendancy over the amphibia, to become in the Mesozoic age the highest and most varied of the existing vertebrates. At that time there were the great land dinosaurs with a length of 80 feet, like Brontosaurus; aquatic forms like Ichthyosaurus and Plesiosaurus, whose mode of evolution from terrestrial to swimming habits was like that of seals and penguins of far later eras. Flying reptiles also evolved, to set an example for the bats of the mammalian class, for both kinds of flying organisms converted their anterior limbs into wings, although in different ways.
During the Triassic and Jurassic periods of the Mesozoic age, the first birds and mammals appeared to follow out their diverging and independent lines of descent. Palaeontology makes it possible to trace the origin and development of many of the different branches that grew out of the mammalian limb from different places and at different times during the Mesozoic and the following age, called the Cenozoic, or age of recent animals. It is unnecessary, however, for us to review more of the details: the main result is obvious; namely, that the appearance of the great classes of vertebrates is in the order of comparative anatomy and embryology. Not only, then, is the fact of evolution rendered trebly sure, but the general order of events is thrice and independently demonstrated to be one and the same. Surely we must see that no reasonable explanation other than evolution can be given for these basic facts and principles.
Turning now to the second division of palaeontological evidence, we come to those groups where abundant materials make it possible to arrange the animals of successive epochs in series that may be remarkably complete. For the reasons specified, the backboned animals provide the richest arrays of these series, and such histories as those of horses and elephants have taken their places in zooelogical science as classics. But even among the invertebrates significant cases may be found. For example, in one restricted locality in Germany the shells of snails belonging to the genus Paludina have been found in superimposed strata in the order of their geological sequence. The ample material shows how the several species altered from age to age by the addition of knobs and ridges to the surface of the shell, until the fossils in the latest rocks are far different from their ancestors in the lowermost levels. Yet the intervening shells fill in the gaps in such a way as to show almost perfectly how the animals worked out their evolutionary history. This example illustrates the nature of many other known series of mollusks and of brachiopods, extending over longer intervals and connecting more widely separated ages like the Secondary and the present period.
Since the doctrine of evolution and its evidences began to occupy the thoughts of the intellectual world at large, no fossil forms have received more attention than the ancient members of the horse tribe. As we have learned, a modern horse is described by comparative anatomy as a one-toed descendant of remote five-toed ancestors. When the hoofed animals of modern times were reviewed as subjects for comparative anatomical study, the odd-toed forms arranged themselves in a series beginning with an animal like an elephant with the full number of five digits on each foot and ending at the opposite extreme with the horse. A reasonable interpretation of these facts was that the animals with fewer toes had evolved from ancestors with five digits, of which the outer ones had progressively disappeared during successive geological periods, while the middle one enlarged correspondingly. The facts provided by palaeontology sustain this contention with absolutely independent testimony. Disregarding some problematical five-toed forms like Phenacodus, the first type of undoubted relationship to modern horses is Hyracotherium, a little animal about three feet long that lived during the Eocene period of the Cenozoic epoch. Its forefeet had four toes each, and its hinder limbs ended with three toes armed with small hoofs, but one of its relatives of the same time has a vestige of another digit on the hind foot. By the geological time mentioned, therefore, the earliest true horses had already lost some of the toes that their progenitors possessed. In the Miocene the extinct species, obviously descended from the Eocene forms, had lost more of their toes; still higher, that is, in the rocks formed during succeeding periods of time, the animals of this division are much larger and each of their feet has only three toes, of which the middle one is the largest while the ones on the sides are small and withdrawn from the ground so as to appear as useless vestiges. To produce modern horses and zebras from these nearer ancestors, few additional changes in the structure of the feet are necessary, for the lateral toes need only to become a little more reduced and the middle one to enlarge slightly to give the one-toed limb of modern types, with its splint-like vestiges still in evidence to show that the ancestor's foot comprised more of these terminal elements. Comparing the animals of successive periods, these and other skeletal structures demonstrate that the ancestry of each group of species is to be found in the animals of the preceding epoch, and that the whole history of horses is one of natural transformation,—in a word, of evolution.
No less interesting in their own way are the remains of other hoofed forms that lead down to the elephants of to-day and to the mammoth and mastodon of relatively recent geologic times. Common sense would lead to the conclusion that a form like a modern tapir was the prototype from which these creatures have arisen, and common sense would lead us to expect that if any fossils of the ancestors of the modern group of elephants occurred at all they would be like tapirs. Thus a fossil of much significance in this connection is Moeritherium, whose remains have been found in the rocks exposed in the Libyan desert, for this creature was practically a tapir, while at the same time its characters of muzzle and tusk mark it as very close to the ancestors of the larger woolly elephants of later geological times, when the trunk had grown considerably and the tusks had become greatly prolonged. Again the fossil sequence confirms the conclusions of comparative anatomy, regarding the mode by which certain modern animals have evolved.
The fossil deer of North America, as well as many other even-toed members of the group of mammalia possessing hoofs, provide the same kind of conclusive evidence. The feature of particular interest in the case of their horns, is a correspondence between the fossil sequence and the order of events in the life-history of existing species,—that is, between the results of palaeontology and of embryology. Horns of the earliest known fossil deer have only two prongs; in the rocks above are remains of deer with additional prongs, and point after point is added as the ancient history of deer is traced upwards through the rocks to modern species. We know that the life-history of a modern species of animals reviews the ancestral record of the species, and what happens during the development of deer can be directly compared with the fossil series. It is a matter of common knowledge that the year-old stag has simple spikes as horns, and that these are shed to be replaced the following year by larger forked horns. Every year the horns are lost and new ones grow out, and become more and more elaborately branched as time goes on, thus giving a series of developmental stages that faithfully repeats the general order of fossil horns. Even Agassiz, who was a believer in special creation and an opponent of evolution, was constrained to point out many other instances, mainly among the invertebrata, where there was a like correspondence between the ontogeny of existing species and their phylogenetic history as revealed by the fossil remains of their ancestors.
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In the last place, we must give more than a passing consideration to some of the extinct types of animals that occupy the position of "links" between groups now widely separated by their divergence in evolution from the same ancestors. Perhaps the most famous example is Archaeopteryx found in a series of slates in Germany. This animal is at once a feathered, flying reptile, and a primitive bird with countless reptilian structures. Its short head possesses lizard-like jaws, all of which bear teeth; its wings comprise five clawed digits; its tail is composed of a long series of joints or vertebrae, bearing large feathers in pairs; its breastbone is flat and like a plate, thus resembling that of reptiles and differing markedly from the great keeled breastbone of modern flying birds, whose large muscles have necessitated the development of the keel for purposes of firm attachment. In brief, this animal was close to the point where reptiles and birds parted company in evolution, and although it was a primitive bird, it is in a true sense a "missing link" between reptiles and the group of modern birds. Other fossil forms like Hesperornis and Ichthyornis, whose remains occur in the strata of a later date, fill in the gap between Archaeopteryx and the birds at the present time, for among other things they possess teeth which indicate their origin from forms like Archaeopteryx, while in other respects they are far nearer the birds of later epochs. That these links are not unique is proved by numerous other examples known to science, such as those which connect amphibia and reptiles, ancient reptiles and primitive mammals, as well as those which come between the different orders of certain vertebrate classes.
In summarizing the foregoing facts, and the larger bodies of evidence that they exemplify, we learn how surely the testimony of the rocks establishes evolution in its own way, how it confirms the law of recapitulation demonstrated by comparative embryology, and how it proves that the greater and smaller divisions of animals have followed the identical order in their evolution that the comparative study of the present day animals has independently described.
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The facts of geographical distribution constitute the fifth division of zooelogy, and an independent class of evidences proving the occurrence of evolution. This department of zooelogy assumed its rightful status only after the other divisions had attained considerable growth. Many naturalists before Darwin and Wallace and Wagner had noticed that animals and plants were by no means evenly distributed over the surface of the globe, but until the doctrine of evolution cleared their vision they did not see the meaning of these facts. As in the case of all the other departments of zooelogy the immediate data themselves are familiar, but because they are so obvious the mind does not look for their interpretation but accepts the facts at their face value. While the phenomena of distribution are no less fascinating to the naturalist, and no less effective in their demonstration of evolution, their comprehensive treatment would demand more space than the whole purpose of the present description of organic evolution would justify. Thus a brief outline only can be given of the salient principles of this subject in order that their bearing upon the problem of species may be indicated.
Even as children we learn many facts of animal distribution; every one knows that lions occur in Africa and not in America, that tigers live in Asia and Malaysia, that the jaguar is an inhabitant of the Brazilian forests, and that the American puma or mountain lion spreads from north to south and from east to west throughout the American continents. The occurrence of differing human races in widely separated localities is no less familiar and striking, for the red man in America, the Zulu in Africa, the Mongol and Malay in their own territories, display the same discontinuity in distribution that is characteristic of all other groups of animals and of plants as well. As our sphere of knowledge increases, we are impressed more and more forcibly by the diversity and unequal extent of the ranges occupied by the members of every one of the varied divisions of the organic world. Another fact which becomes significant only when science calls our attention to it is the absence from a land like Australia of higher mammals such as the rabbit of Europe. The hypothesis of special creation cannot explain this absence on the assumption that the rabbit is unsuited to the conditions obtaining in the country named, for when the species was introduced into Australia by man, it developed and spread with marvelous rapidity and destructive effect. It may seem impossible that facts like these could possess an evolutionary significance, but they are actual examples of the great mass of data brought together by the naturalists who have seen in them something to be interpreted, and who have sought and found an explanation in the formularies of science.
The general principles of distribution appear with greatest clearness when an examination is made of the animals and plants of isolated regions like islands. The Galapagos Islands constitute a group that has figured largely in the literature of the subject, partly because Darwin himself was so impressed by what he found there in the course of his famous voyage around the world in the "Beagle." They form a cluster on the Equator about six hundred miles west of the nearest point of the neighboring coast of South America. Although the lizards and birds that live in the group differ somewhat among themselves as one passes from island to island, on the whole they are most like the species of the corresponding classes inhabiting South America. Why should this be so? On the hypothesis of special creation there is no reason why they should not be more like the species of Africa or Australia than like those of the nearest body of the mainland. The explanation given by evolution is clear, simple, and reasonable. It is that the characteristic island forms are the descendants of immigrants which in greatest probability would be wanderers from the neighboring continent and not from far distant lands. Reaching the isolated area in question the natural factors of evolution would lead their offspring of later generations to vary from the original parental types, and so the peculiar Galapagos species would come into being. The fact that the organisms living on the various islands of this group differ somewhat in lesser details adds further justification for the evolutionary interpretation, because it is not probable that all the islands would be populated at the same time by similar stragglers from the mainland. The first settlers in one place would send out colonies to others, where independent evolution would result in the appearance of minor differences peculiar to the single island. In this manner science interprets the general agreement between the animals of the Azores Islands and the fauna of the northwestern part of Africa, the nearest body of land, from which it would be most natural for the ancestors of the island fauna to come.
The land-snails inhabiting the various groups of islands scattered throughout the vast extent of the Pacific Ocean provide the richest and most ideal material for the demonstration of the principles of geographical distribution. In the Hawaiian Islands snails of the family of Achatinellidae occur in great abundance, and like the lizards of the Galapagos Islands different species occur on the different members of the group. Within the confines of one and the same island, they vary from valley to valley, and the correlation between their isolation in geographical respects and specific differences on the other hand, first pointed out by Gulick, makes this tribe of animals classical material. In Polynesia and Melanesia are found close relatives of the Achatinellidae, namely, the Partulae, which are thus in relative proximity to the Achatinellidae and not on the other side of the world. Furthermore, the Partulae are not alike in all of the groups of Polynesia where they occur; the species of the Society Islands are absolutely distinct from those of the Marquesas, Tonga, Samoan, and Solomon Islands, although they agree closely in the basic characters that justify their reference to a single genus. The geological evidence tells us that these islands were once the peaks of mountain ranges rising from a Pacific continent which has since subsided to such an extent that the mountain tops have become separate islands. Thus the resemblances between Hawaiian and Polynesian snails, and the closer similarities exhibited by the species of the various groups of Polynesia, are intelligible as the marks of a common ancestry in a widespread continental stock, while the observed differences show the extent of subsequent evolution along independent lines followed out after the isolation of the now separated islands. The principle may be worked out in even greater detail, for it appears that within the limits of one group diverse forms occupy different islands, evolved in different ways in their own neighborhoods; while in one and the same island, the populations of the different valleys show marked effects of divergence in later evolution, precisely as in the case of the classic Achatinellidae of the Hawaiian Islands.
The broad and consistent principle underlying these and related facts is this: there is a general correspondence between the differences displayed by the organisms of two regions and the degree of isolation or proximity of these two areas. Thus the disconnected but neighboring areas of the Galapagos Islands and South America support species that resemble each other closely, for the reasons given before; long isolated areas like Australia and its surroundings possess peculiar creatures like the egg-laying mammals, and all of the pouched animals or marsupials with only one or two exceptions like our own American opossum,—a correlation between a geological and geographical discontinuity on the one hand and a peculiarity on the other that reinforces our confidence in the faunal evolutionary interpretation of the facts of distribution.
It is true that the various classes of animals do not always appear with coextensive ranges. The barriers between two groups of related species will not be the same in all cases. A range like the Rocky Mountains will keep fresh-water fish apart, while birds and mammals can get across somewhere at some time. All these things must be taken into account in analyzing the phenomena of distribution, and many factors must be given due attention; but in all cases the reasons for the particular state of affairs in geographical and biological respects possess an evolutionary significance.
Having then all the facts of animal natural history at his disposal, and the uniform principles in each body of fact that demonstrate evolution, it is small wonder that the evolutionist seems to dogmatize when he asserts that descent with adaptive and divergent modification is true for all species of living things. The case is complete as it stands to-day, while it is even more significant that every new discovery falls into line with what is already known, and takes its natural place in the all-inclusive doctrine of organic evolution. Because this explanation of the characteristics of the living world is more reasonable than any other, science teaches that it is true.
EVOLUTION AS A NATURAL PROCESS
The purpose of the discussions up to this point has been to present the reasons drawn from the principal classes of zooelogical facts for believing that living things have transformed naturally to become what they now are. Even if it were possible to make an exhaustive analysis of all of the known phenomena of animal structure, development, and fossil succession, the complete bodies of knowledge could not make the evolutionary explanation more real and evident than it is shown to be by the simple facts and principles selected to constitute the foregoing outline. We have dealt solely with the evidences as to the fact of evolution; and now, having assured ourselves that it is worth while to so do, we may turn to the intelligible and reasonable evidence found by science which proves that the familiar and everyday "forces" of nature are competent to bring about evolution if they have operated in the past as they do to-day. Investigation has brought to light many of the subsidiary elements of the whole process, and these are so real and obvious that they are simply taken for granted without a suspicion on our part of their power until science directs our attention to them.
For one reason or another, those who take up this subject for the first time find it difficult to banish from their minds the idea that evolution, even if it ever took place, has been ended. They think it futile to expect that a scrutiny of to-day's order can possibly find influences powerful enough to have any share in the marvelous process of past evolution demonstrated by science. The naturalists of a century ago held a similar opinion regarding the earth, viewing it as an immutable and unchanged product of supernatural creation, until Lyell led them to see that the world is a plastic mass slowly altering in countless ways. It is no more true that living things have ceased to evolve than that mountains and rivers and glaciers are fixed in their final forms; they may seem everlasting and permanent only because a human life is so brief in comparison with their full histories. Like the development of a continent as science describes it, the origin of a new species by evolution, its rise, culmination, and final extinction may demand thousands of years; so that an onlooker who is himself only a conscious atom of the turbulent stream of evolving organic life does not live long enough to observe more than a small fraction of the whole process. Therefore living species seem unchanged and unchangeable until a conviction that evolution is true, and a knowledge of the method of science by which this conviction is borne upon one, guide the student onwards in the further search for the efficient causes of the process.
The biologist employs the identical methods used by the geologist in working out the past history of the earth's crust. The latter observes the forces at work to-day, and compares the new layers of rock now being formed with the strata of deeper levels; these are so much alike that he is led to regard the constructive influences of the past as identical with those he can now watch at work. Similarly the biologist must first learn, as we have done, the principles of animal construction and development, and of other classes of zooelogical facts, and then he must turn his attention from the dead object of laboratory analysis to the workings of organic machines. The way an organism lives its life in dynamic relations to the varied conditions of existence, as well as the mutual physiological relations of the manifold parts of a single organism, reveal certain definite natural forces at work. Therefore his next task is to compare the results accomplished by these factors in the brief time they may be seen in operation with the products of the whole process of organic evolution, to learn, like the geologist in his sphere, that the present-day natural forces are able to do what reason says they have done in the past.