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Darwin and Modern Science (1909)

Edited by A.C. Seward


XI. THE PALAEONTOLOGICAL RECORD.

I. ANIMALS.

By W.B. SCOTT.
Professor of Geology in the University of Princeton, U.S.A.

S
o no branch of science did the publication of "The Origin of Species" prove to be a more vivifying and transforming influence than to Palaeontology. This science had suffered, and to some extent, still suffers from its rather anomalous position between geology and biology, each of which makes claim to its territory, and it was held in strict bondage to the Linnean and Cuvierian dogma that species were immutable entities. There is, however, reason to maintain that this strict bondage to a dogma now abandoned, was not without its good side, and served the purpose of keeping the infant science in leading-strings until it was able to walk alone, and preventing a flood of premature generalisations and speculations.

As Zittel has said: "Two directions were from the first apparent in palaeontological research—a stratigraphical and a biological. Stratigraphers wished from palaeontology mainly confirmation regarding the true order or relative age of zones of rock-deposits in the field. Biologists had, theoretically at least, the more genuine interest in fossil organisms as individual forms of life." (Zittel, "History of Geology and Palaeontology", page 363, London, 1901.) The geological or stratigraphical direction of the science was given by the work of William Smith, "the father of historical geology," in the closing decade of the eighteenth century. Smith was the first to make a systematic use of fossils in determining the order of succession of the rocks which make up the accessible crust of the earth, and this use has continued, without essential change, to the present day. It is true that the theory of evolution has greatly modified our conceptions concerning the introduction of new species and the manner in which palaeontological data are to be interpreted in terms of stratigraphy, but, broadly speaking, the method remains fundamentally the same as that introduced by Smith.

The biological direction of palaeontology was due to Cuvier and his associates, who first showed that fossils were not merely varieties of existing organisms, but belonged to extinct species and genera, an altogether revolutionary conception, which startled the scientific world. Cuvier made careful studies, especially of fossil vertebrates, from the standpoint of zoology and was thus the founder of palaeontology as a biological science. His great work on "Ossements Fossiles" (Paris, 1821) has never been surpassed as a masterpiece of the comparative method of anatomical investigation, and has furnished to the palaeontologist the indispensable implements of research.

On the other hand, Cuvier's theoretical views regarding the history of the earth and its successive faunas and floras are such as no one believes to- day. He held that the earth had been repeatedly devastated by great cataclysms, which destroyed every living thing, necessitating an entirely new creation, thus regarding the geological periods as sharply demarcated and strictly contemporaneous for the whole earth, and each species of animal and plant as confined to a single period. Cuvier's immense authority and his commanding personality dominated scientific thought for more than a generation and marked out the line which the development of palaeontology was to follow. The work was enthusiastically taken up by many very able men in the various European countries and in the United States, but, controlled as it was by the belief in the fixity of species, it remained almost entirely descriptive and consisted in the description and classification of the different groups of fossil organisms. As already intimated, this narrowness of view had its compensations, for it deferred generalisations until some adequate foundations for these had been laid.

Dominant as it was, Cuvier's authority was slowly undermined by the progress of knowledge and the way was prepared for the introduction of more rational conceptions. The theory of "Catastrophism" was attacked by several geologists, most effectively by Sir Charles Lyell, who greatly amplified the principles enunciated by Hutton and Playfair in the preceding century, and inaugurated a new era in geology. Lyell's uniformitarian views of the earth's history and of the agencies which had wrought its changes, had undoubted effect in educating men's minds for the acceptance of essentially similar views regarding the organic world. In palaeontology too the doctrine of the immutability of species, though vehemently maintained and reasserted, was gradually weakening. In reviewing long series of fossils, relations were observed which pointed to genetic connections and yet were interpreted as purely ideal. Agassiz, for example, who never accepted the evolutionary theory, drew attention to facts which could be satisfactorily interpreted only in terms of that theory. Among the fossils he indicated "progressive," "synthetic," "prophetic," and "embryonic" types, and pointed out the parallelism which obtains between the geological succession of ancient animals and the ontogenetic development of recent forms. In Darwin's words: "This view accords admirably well with our theory." ("Origin of Species" (6th edition), page 310.) Of similar import were Owen's views on "generalised types" and "archetypes."

The appearance of "The Origin of Species" in 1859 revolutionised all the biological sciences. From the very nature of the case, Darwin was compelled to give careful consideration to the palaeontological evidence; indeed, it was the palaeontology and modern distribution of animals in South America which first led him to reflect upon the great problem. In his own words: "I had been deeply impressed by discovering in the Pampean formation great fossil animals covered with armour like that on the existing armadillos; secondly, by the manner in which closely allied animals replace one another in proceeding southward over the Continent; and thirdly, by the South American character of most of the productions of the Galapagos archipelago, and more especially by the manner in which they differ slightly on each island of the group." ("Life and Letters of Charles Darwin", I. page 82.) In the famous tenth and eleventh chapters of the "Origin", the palaeontological evidence is examined at length and the imperfection of the geological record is strongly emphasised. The conclusion is reached, that, in view of this extreme imperfection, palaeontology could not reasonably be expected to yield complete and convincing proof of the evolutionary theory. "I look at the geological record as a history of the world imperfectly kept, and written in a changing dialect; of this history we possess the last volume alone, relating only to two or three countries. Of this volume, only here and there a short chapter has been preserved; and of each page, only here and there a few lines." ("Origin of Species", page 289.) Yet, aside from these inevitable difficulties, he concludes, that "the other great leading facts in palaeontology agree admirably with the theory of descent with modification through variation and natural selection." (Ibid. page 313.)

Darwin's theory gave an entirely new significance and importance to palaeontology. Cuvier's conception of the science had been a limited, though a lofty one. "How glorious it would be if we could arrange the organised products of the universe in their chronological order!...The chronological succession of organised forms, the exact determination of those types which appeared first, the simultaneous origin of certain species and their gradual decay, would perhaps teach us as much about the mysteries of organisation as we can possibly learn through experiments with living organisms." (Zittel op. cit. page 140.) This, however, was rather the expression of a hope for the distant future than an account of what was attainable, and in practice the science remained almost purely descriptive, until Darwin gave it a new standpoint, new problems and an altogether fresh interest and charm. The revolution thus accomplished is comparable only to that produced by the Copernican astronomy.

From the first it was obvious that one of the most searching tests of the evolutionary theory would be given by the advance of palaeontological discovery. However imperfect the geological record might be, its ascertained facts would necessarily be consistent, under any reasonable interpretation, with the demands of a true theory; otherwise the theory would eventually be overwhelmed by the mass of irreconcilable data. A very great stimulus was thus given to geological investigation and to the exploration of new lands. In the last forty years, the examination of North and South America, of Africa and Asia has brought to light many chapters in the history of life, which are astonishingly full and complete. The flood of new material continues to accumulate at such a rate that it is impossible to keep abreast of it, and the very wealth of the collections is a source of difficulty and embarrassment. In modern palaeontology phylogenetic questions and problems occupy a foremost place and, as a result of the labours of many eminent investigators in many lands, it may be said that this science has proved to be one of the most solid supports of Darwin's theory. True, there are very many unsolved problems, and the discouraged worker is often tempted to believe that the fossils raise more questions than they answer. Yet, on the other hand, the whole trend of the evidence is so strongly in favour of the evolutionary doctrine, that no other interpretation seems at all rational.

To present any adequate account of the palaeontological record from the evolutionary standpoint, would require a large volume and a singularly unequal, broken and disjointed history it would be. Here the record is scanty, interrupted, even unintelligible, while there it is crowded with embarrassing wealth of material, but too often these full chapters are separated by such stretches of unrecorded time, that it is difficult to connect them. It will be more profitable to present a few illustrative examples than to attempt an outline of the whole history.

At the outset, the reader should be cautioned not to expect too much, for the task of determining phylogenies fairly bristles with difficulties and encounters many unanswered questions. Even when the evidence seems to be as copious and as complete as could be wished, different observers will put different interpretations upon it, as in the notorious case of the Steinheim shells. (In the Miocene beds of Steinheim, Wurtemberg, occur countless fresh-water shells, which show numerous lines of modification, but these have been very differently interpreted by different writers.) The ludicrous discrepances which often appear between the phylogenetic "trees" of various writers have cast an undue discredit upon the science and have led many zoologists to ignore palaeontology altogether as unworthy of serious attention. One principal cause of these discrepant and often contradictory results is our ignorance concerning the exact modes of developmental change. What one writer postulates as almost axiomatic, another will reject as impossible and absurd. Few will be found to agree as to how far a given resemblance is offset by a given unlikeness, and so long as the question is one of weighing evidence and balancing probabilities, complete harmony is not to be looked for. These formidable difficulties confront us even in attempting to work out from abundant material a brief chapter in the phylogenetic history of some small and clearly limited group, and they become disproportionately greater, when we extend our view over vast periods of time and undertake to determine the mutual relationships of classes and types. If the evidence were complete and available, we should hardly be able to unravel its infinite complexity, or to find a clue through the mazes of the labyrinth. "Our ideas of the course of descent must of necessity be diagrammatic." (D.H. Scott, "Studies in Fossil Botany", page 524. London, 1900.)

Some of the most complete and convincing examples of descent with modification are to be found among the mammals, and nowhere more abundantly than in North America, where the series of continental formations, running through the whole Tertiary period, is remarkably full. Most of these formations contain a marvellous wealth of mammalian remains and in an unusual state of preservation. The oldest Eocene (Paleocene) has yielded a mammalian fauna which is still of prevailingly Mesozoic character, and contains but few forms which can be regarded as ancestral to those of later times. The succeeding fauna of the lower Eocene proper (Wasatch stage) is radically different and, while a few forms continue over from the Paleocene, the majority are evidently recent immigrants from some region not yet identified. From the Wasatch onward, the development of many phyla may be traced in almost unbroken continuity, though from time to time the record is somewhat obscured by migrations from the Old World and South America. As a rule, however, it is easy to distinguish between the immigrant and the indigenous elements of the fauna.

From their gregarious habits and individual abundance, the history of many hoofed animals is preserved with especial clearness. So well known as to have become a commonplace, is the phylogeny of the horses, which, contrary to all that would have been expected, ran the greater part of its course in North America. So far as it has yet been traced, the line begins in the lower Eocene with the genus Eohippus, a little creature not much larger than a cat, which has a short neck, relatively short limbs, and in particular, short feet, with four functional digits and a splint-like rudiment in the fore-foot, three functional digits and a rudiment in the hind-foot. The forearm bones (ulna and radius) are complete and separate, as are also the bones of the lower leg (fibula and tibia). The skull has a short face, with the orbit, or eye-socket, incompletely enclosed with bone, and the brain-case is slender and of small capacity. The teeth are short- crowned, the incisors without "mark," or enamel pit, on the cutting edge; the premolars are all smaller and simpler than the molars. The pattern of the upper molars is so entirely different from that seen in the modern horses that, without the intermediate connecting steps, no one would have ventured to derive the later from the earlier plan. This pattern is quadritubercular, with four principal, conical cusps arranged in two transverse pairs, forming a square, and two minute cuspules between each transverse pair, a tooth which is much more pig-like than horse-like. In the lower molars the cusps have already united to form two crescents, one behind the other, forming a pattern which is extremely common in the early representatives of many different families, both of the Perissodactyla and the Artiodactyla. In spite of the manifold differences in all parts of the skeleton between Eohippus and the recent horses, the former has stamped upon it an equine character which is unmistakable, though it can hardly be expressed in words.

Each one of the different Eocene and Oligocene horizons has its characteristic genus of horses, showing a slow, steady progress in a definite direction, all parts of the structure participating in the advance. It is not necessary to follow each of these successive steps of change, but it should be emphasised that the changes are gradual and uninterrupted. The genus Mesohippus, of the middle Oligocene, may be selected as a kind of half-way stage in the long progression. Comparing Mesohippus with Eohippus, we observe that the former is much larger, some species attaining the size of a sheep, and has a relatively longer neck, longer limbs and much more elongate feet, which are tridactyl, and the middle toe is so enlarged that it bears most of the weight, while the lateral digits are very much more slender. The fore-arm bones have begun to co-ossify and the ulna is greatly reduced, while the fibula, though still complete, is hardly more than a thread of bone. The skull has a longer face and a nearly enclosed orbit, and the brain-case is fuller and more capacious, the internal cast of which shows that the brain was richly convoluted. The teeth are still very short-crowned, but the upper incisors plainly show the beginning of the "mark"; the premolars have assumed the molar form, and the upper molars, though plainly derived from those of Eohippus, have made a long stride toward the horse pattern, in that the separate cusps have united to form a continuous outer wall and two transverse crests.

In the lower Miocene the interesting genus Desmatippus shows a further advance in the development of the teeth, which are beginning to assume the long-crowned shape, delaying the formation of roots; a thin layer of cement covers the crowns, and the transverse crests of the upper grinding teeth display an incipient degree of their modern complexity. This tooth-pattern is strictly intermediate between the recent type and the ancient type seen in Mesohippus and its predecessors. The upper Miocene genera, Protohippus and Hipparion are, to all intents and purposes, modern in character, but their smaller size, tridactyl feet and somewhat shorter-crowned teeth are reminiscences of their ancestry.

From time to time, when a land-connection between North America and Eurasia was established, some of the successive equine genera migrated to the Old World, but they do not seem to have gained a permanent footing there until the end of the Miocene or beginning of the Pliocene, eventually diversifying into the horses, asses, and zebras of Africa, Asia and Europe. At about the same period, the family extended its range to South America and there gave rise to a number of species and genera, some of them extremely peculiar. For some unknown reason, all the horse tribe had become extinct in the western hemisphere before the European discovery, but not until after the native race of man had peopled the continents.

In addition to the main stem of equine descent, briefly considered in the foregoing paragraphs, several side-branches were given off at successive levels of the stem. Most of these branches were short-lived, but some of them flourished for a considerable period and ramified into many species.

Apparently related to the horses and derived from the same root-stock is the family of the Palaeotheres, confined to the Eocene and Oligocene of Europe, dying out without descendants. In the earlier attempts to work out the history of the horses, as in the famous essay of Kowalevsky ("Sur l'Anchitherium aurelianense Cuv. et sur l'histoire paleontologique des Chevaux", "Mem. de l'Acad. Imp. des Sc. de St Petersbourg", XX. no. 5, 1873.), the Palaeotheres were placed in the direct line, because the number of adequately known Eocene mammals was then so small, that Cuvier's types were forced into various incongruous positions, to serve as ancestors for unrelated series.

The American family of the Titanotheres may also be distantly related to the horses, but passed through an entirely different course of development. From the lower Eocene to the lower sub-stage of the middle Oligocene the series is complete, beginning with small and rather lightly built animals. Gradually the stature and massiveness increase, a transverse pair of nasal horns make their appearance and, as these increase in size, the canine tusks and incisors diminish correspondingly. Already in the oldest known genus the number of digits had been reduced to four in the fore-foot and three in the hind, but there the reduction stops, for the increasing body- weight made necessary the development of broad and heavy feet. The final members of the series comprise only large, almost elephantine animals, with immensely developed and very various nasal horns, huge and massive heads, and altogether a grotesque appearance. The growth of the brain did not at all keep pace with the increase of the head and body, and the ludicrously small brain may will have been one of the factors which determined the startlingly sudden disappearance and extinction of the group.

Less completely known, but of unusual interest, is the genealogy of the rhinoceros family, which probably, though not certainly, was likewise of American origin. The group in North America at least, comprised three divisions, or sub-families, of very different proportions, appearance and habits, representing three divergent lines from the same stem. Though the relationship between the three lines seems hardly open to question, yet the form ancestral to all of them has not yet been identified. This is because of our still very incomplete knowledge of several perissodactyl genera of the Eocene, any one of which may eventually prove to be the ancestor sought for.

The first sub-family is the entirely extinct group of Hyracodonts, which may be traced in successive modifications through the upper Eocene, lower and middle Oligocene, then disappearing altogether. As yet, the hyracodonts have been found only in North America, and the last genus of the series, Hyracodon, was a cursorial animal. Very briefly stated, the modifications consist in a gradual increase in size, with greater slenderness of proportions, accompanied by elongation of the neck, limbs, and feet, which become tridactyl and very narrow. The grinding teeth have assumed the rhinoceros-like pattern and the premolars resemble the molars in form; on the other hand, the front teeth, incisors and canines, have become very small and are useless as weapons. As the animal had no horns, it was quite defenceless and must have found its safety in its swift running, for Hyracodon displays many superficial resemblances to the contemporary Oligocene horses, and was evidently adapted for speed. It may well have been the competition of the horses which led to the extinction of these cursorial rhinoceroses.

The second sub-family, that of the Amynodonts, followed a totally different course of development, becoming short-legged and short-footed, massive animals, the proportions of which suggest aquatic habits; they retained four digits in the front foot. The animal was well provided with weapons in the large canine tusks, but was without horns. Some members of this group extended their range to the Old World, but they all died out in the middle Oligocene, leaving no successors.

The sub-family of the true rhinoceroses cannot yet be certainly traced farther back than to the base of the middle Oligocene, though some fragmentary remains found in the lower Oligocene are probably also referable to it. The most ancient and most primitive member of this series yet discovered, the genus Trigonias, is unmistakably a rhinoceros, yet much less massive, having more the proportions of a tapir; it had four toes in the front foot, three in the hind, and had a full complement of teeth, except for the lower canines, though the upper canines are about to disappear, and the peculiar modification of the incisors, characteristic of the true rhinoceroses, is already apparent; the skull is hornless. Representatives of this sub-family continue through the Oligocene and Miocene of North America, becoming rare and localised in the Pliocene and then disappearing altogether. In the Old World, on the other hand, where the line appeared almost as early as it did in America, this group underwent a great expansion and ramification, giving rise not only to the Asiatic and African forms, but also to several extinct series.

Turning now to the Artiodactyla, we find still another group of mammals, that of the camels and llamas, which has long vanished from North America, yet took its rise and ran the greater part of its course in that continent. From the lower Eocene onward the history of this series is substantially complete, though much remains to be learned concerning the earlier members of the family. The story is very like that of the horses, to which in many respects it runs curiously parallel. Beginning with very small, five-toed animals, we observe in the successive genera a gradual transformation in all parts of the skeleton, an elongation of the neck, limbs and feet, a reduction of the digits from five to two, and eventually the coalescence of the remaining two digits into a "cannon-bone." The grinding teeth, by equally gradual steps, take on the ruminant pattern. In the upper Miocene the line divides into the two branches of the camels and llamas, the former migrating to Eurasia and the latter to South America, though representatives of both lines persisted in North America until a very late period. Interesting side-branches of this line have also been found, one of which ended in the upper Miocene in animals which had almost the proportions of the giraffes and must have resembled them in appearance.

The American Tertiary has yielded several other groups of ruminant-like animals, some of which form beautifully complete evolutionary series, but space forbids more than this passing mention of them.

It was in Europe that the Artiodactyla had their principal development, and the upper Eocene, Oligocene and Miocene are crowded with such an overwhelming number and variety of forms that it is hardly possible to marshal them in orderly array and determine their mutual relationships. Yet in this chaotic exuberance of life, certain important facts stand out clearly, among these none is of greater interest and importance than the genealogy of the true Ruminants, or Pecora, which may be traced from the upper Eocene onward. The steps of modification and change are very similar to those through which the camel phylum passed in North America, but it is instructive to note that, despite their many resemblances, the two series can be connected only in their far distant beginnings. The pecoran stock became vastly more expanded and diversified than did the camel line and was evidently more plastic and adaptable, spreading eventually over all the continents except Australia, and forming to-day one of the dominant types of mammals, while the camels are on the decline and not far from extinction. The Pecora successively ramified into the deer, antelopes, sheep, goats and oxen, and did not reach North America till the Miocene, when they were already far advanced in specialisation. To this invasion of the Pecora, or true ruminants, it seems probable that the decline and eventual disappearance of the camels is to be ascribed.

Recent discoveries in Egypt have thrown much light upon a problem which long baffled the palaeontologist, namely, the origin of the elephants. (C.W. Andrews, "On the Evolution of the Proboscidea", "Phil. Trans. Roy. Soc." London, Vol. 196, 1904, page 99.) Early representatives of this order, Mastodons, had appeared almost simultaneously (in the geological sense of that word) in the upper Miocene of Europe and North America, but in neither continent was any more ancient type known which could plausibly be regarded as ancestral to them. Evidently, these problematical animals had reached the northern continents by migrating from some other region, but no one could say where that region lay. The Eocene and Oligocene beds of the Fayoum show us that the region sought for is Africa, and that the elephants form just such a series of gradual modifications as we have found among other hoofed animals. The later steps of the transformation, by which the mastodons lost their lower tusks, and their relatively small and simple grinding teeth acquired the great size and highly complex structure of the true elephants, may be followed in the uppermost Miocene and Pliocene fossils of India and southern Europe.

Egypt has also of late furnished some very welcome material which contributes to the solution of another unsolved problem which had quite eluded research, the origin of the whales. The toothed-whales may be traced back in several more or less parallel lines as far as the lower Miocene, but their predecessors in the Oligocene are still so incompletely known that safe conclusions can hardly be drawn from them. In the middle Eocene of Egypt, however, has been found a small, whale-like animal (Protocetus), which shows what the ancestral toothed-whale was like, and at the same time seems to connect these thoroughly marine mammals with land- animals. Though already entirely adapted to an aquatic mode of life, the teeth, skull and backbone of Protocetus display so many differences from those of the later whales and so many approximations to those of primitive, carnivorous land-mammals, as, in a large degree, to bridge over the gap between the two groups. Thus one of the most puzzling of palaeontological questions is in a fair way to receive a satisfactory answer. The origin of the whalebone-whales and their relations to the toothed-whales cannot yet be determined, since the necessary fossils have not been discovered.

Among the carnivorous mammals, phylogenetic series are not so clear and distinct as among the hoofed animals, chiefly because the carnivores are individually much less abundant, and well-preserved skeletons are among the prizes of the collector. Nevertheless, much has already been learned concerning the mutual relations of the carnivorous families, and several phylogenetic series, notably that of the dogs, are quite complete. It has been made extremely probable that the primitive dogs of the Eocene represent the central stock, from which nearly or quite all the other families branched off, though the origin and descent of the cats have not yet been determined.

It should be clearly understood that the foregoing account of mammalian descent is merely a selection of a few representative cases and might be almost indefinitely extended. Nothing has been said, for example, of the wonderful museum of ancient mammalian life which is entombed in the rocks of South America, especially of Patagonia, and which opens a world so entirely different from that of the northern continents, yet exemplifying the same laws of "descent with modification." Very beautiful phylogenetic series have already been established among these most interesting and marvellously preserved fossils, but lack of space forbids a consideration of them.

The origin of the mammalia, as a class, offers a problem of which palaeontology can as yet present no definitive solution. Many morphologists regard the early amphibia as the ancestral group from which the mammals were derived, while most palaeontologists believe that the mammals are descended from the reptiles. The most ancient known mammals, those from the upper Triassic of Europe and North America, are so extremely rare and so very imperfectly known, that they give little help in determining the descent of the class, but, on the other hand, certain reptilian orders of the Permian period, especially well represented in South Africa, display so many and such close approximations to mammalian structure, as strongly to suggest a genetic relationship. It is difficult to believe that all those likenesses should have been independently acquired and are without phylogenetic significance.

Birds are comparatively rare as fossils and we should therefore look in vain among them for any such long and closely knit series as the mammals display in abundance. Nevertheless, a few extremely fortunate discoveries have made it practically certain that birds are descended from reptiles, of which they represent a highly specialised branch. The most ancient representative of this class is the extraordinary genus Archaeopteryx from the upper Jurassic of Bavaria, which, though an unmistakable bird, retains so many reptilian structures and characteristics as to make its derivation plain. Not to linger over anatomical minutiae, it may suffice to mention the absence of a horny beak, which is replaced by numerous true teeth, and the long lizard-like tail, which is made up of numerous distinct vertebrae, each with a pair of quill-like feathers attached to it. Birds with teeth are also found in the Cretaceous, though in most other respects the birds of that period had attained a substantially modern structure. Concerning the interrelations of the various orders and families of birds, palaeontology has as yet little to tell us.

The life of the Mesozoic era was characterised by an astonishing number and variety of reptiles, which were adapted to every mode of life, and dominated the air, the sea and the land, and many of which were of colossal proportions. Owing to the conditions of preservation which obtained during the Mesozoic period, the history of the reptiles is a broken and interrupted one, so that we can make out many short series, rather than any one of considerable length. While the relations of several reptilian orders can be satisfactorily determined, others still baffle us entirely, making their first known appearance in a fully differentiated state. We can trace the descent of the sea-dragons, the Ichthyosaurs and Plesiosaurs, from terrestrial ancestors, but the most ancient turtles yet discovered show us no closer approximation to any other order than do the recent turtles; and the oldest known Pterosaurs, the flying dragons of the Jurassic, are already fully differentiated. There is, however, no ground for discouragement in this, for the progress of discovery has been so rapid of late years, and our knowledge of Mesozoic life has increased with such leaps and bounds, that there is every reason to expect a solution of many of the outstanding problems in the near future.

Passing over the lower vertebrates, for lack of space to give them any adequate consideration, we may briefly take up the record of invertebrate life. From the overwhelming mass of material it is difficult to make a representative selection and even more difficult to state the facts intelligibly without the use of unduly technical language and without the aid of illustrations.

Several groups of the Mollusca, or shell-fish, yield very full and convincing evidence of their descent from earlier and simpler forms, and of these none is of greater interest than the Ammonites, an extinct order of the cephalopoda. The nearest living ally of the ammonites is the pearly nautilus, the other existing cephalopods, such as the squids, cuttle-fish, octopus, etc., are much more distantly related. Like the nautilus, the ammonites all possess a coiled and chambered shell, but their especial characteristic is the complexity of the "sutures." By sutures is meant the edges of the transverse partitions, or septa, where these join the shell- wall, and their complexity in the fully developed genera is extraordinary, forming patterns like the most elaborate oak-leaf embroidery, while in the nautiloids the sutures form simple curves. In the rocks of the Mesozoic era, wherever conditions of preservation are favourable, these beautiful shells are stored in countless multitudes, of an incredible variety of form, size and ornamentation, as is shown by the fact that nearly 5000 species have already been described. The ammonites are particularly well adapted for phylogenetic studies, because, by removing the successive whorls of the coiled shell, the individual development may be followed back in inverse order, to the microscopic "protoconch," or embryonic shell, which lies concealed in the middle of the coil. Thus the valuable aid of embryology is obtained in determining relationships.

The descent of the ammonites, taken as a group, is simple and clear; they arose as a branch of the nautiloids in the lower Devonian, the shells known as goniatites having zigzag, angulated sutures. Late in the succeeding Carboniferous period appear shells with a truly ammonoid complexity of sutures, and in the Permian their number and variety cause them to form a striking element of the marine faunas. It is in the Mesozoic era, however, that these shells attain their full development; increasing enormously in the Triassic, they culminate in the Jurassic in the number of families, genera and species, in the complexity of the sutures, and in the variety of shell-ornamentation. A slow decline begins in the Cretaceous, ending in the complete extinction of the whole group at the end of that period. As a final phase in the history of the ammonites, there appear many so-called "abnormal" genera, in which the shell is irregularly coiled, or more or less uncoiled, in some forms becoming actually straight. It is interesting to observe that some of these genera are not natural groups, but are "polyphyletic," i.e. are each derived from several distinct ancestral genera, which have undergone a similar kind of degeneration.

In the huge assembly of ammonites it is not yet possible to arrange all the forms in a truly natural classification, which shall express the various interrelations of the genera, yet several beautiful series have already been determined. In these series the individual development of the later general shows transitory stages which are permanent in antecedent genera. To give a mere catalogue of names without figures would not make these series more intelligible.

The Brachiopoda, or "lamp-shells," are a phylum of which comparatively few survive to the present day; their shells have a superficial likeness to those of the bivalved Mollusca, but are not homologous with the latter, and the phylum is really very distinct from the molluscs. While greatly reduced now, these animals were incredibly abundant throughout the Palaeozoic era, great masses of limestone being often composed almost exclusively of their shells, and their variety is in keeping with their individual abundance. As in the case of the ammonites, the problem is to arrange this great multitude of forms in an orderly array that shall express the ramifications of the group according to a genetic system. For many brachiopods, both recent and fossil, the individual development, or ontogeny, has been worked out and has proved to be of great assistance in the problems of classification and phylogeny. Already very encouraging progress has been made in the solution of these problems. All brachiopods form first a tiny, embryonic shell, called the protegulum, which is believed to represent the ancestral form of the whole group, and in the more advanced genera the developmental stages clearly indicate the ancestral genera of the series, the succession of adult forms in time corresponding to the order of the ontogenetic stages. The transformation of the delicate calcareous supports of the arms, often exquisitely preserved, are extremely interesting. Many of the Palaeozoic genera had these supports coiled like a pair of spiral springs, and it has been shown that these genera were derived from types in which the supports were simply shelly loops.

The long extinct class of crustacea known as the Trilobites are likewise very favourable subjects for phylogenetic studies. So far as the known record can inform us, the trilobites are exclusively Palaeozoic in distribution, but their course must have begun long before that era, as is shown by the number of distinct types among the genera of the lower Cambrian. The group reached the acme of abundance and relative importance in the Cambrian and Ordovician; then followed a long, slow decline, ending in complete and final disappearance before the end of the Permian. The newly-hatched and tiny trilobite larva, known as the protaspis, is very near to the primitive larval form of all the crustacea. By the aid of the correlated ontogenetic stages and the succession of the adult forms in the rocks, many phylogenetic series have been established and a basis for the natural arrangement of the whole class has been laid.

Very instructive series may also be observed among the Echinoderms and, what is very rare, we are able in this sub-kingdom to demonstrate the derivation of one class from another. Indeed, there is much reason to believe that the extinct class Cystidea of the Cambrian is the ancestral group, from which all the other Echinoderms, star-fishes, brittle-stars, sea-urchins, feather-stars, etc., are descended.

The foregoing sketch of the palaeontological record is, of necessity, extremely meagre, and does not represent even an outline of the evidence, but merely a few illustrative examples, selected almost at random from an immense body of material. However, it will perhaps suffice to show that the geological record is not so hopelessly incomplete as Darwin believed it to be. Since "The Origin of Species" was written, our knowledge of that record has been enormously extended and we now possess, no complete volumes, it is true, but some remarkably full and illuminating chapters. The main significance of the whole lies in the fact, that just in proportion to the completeness of the record is the unequivocal character of its testimony to the truth of the evolutionary theory.

The test of a true, as distinguished from a false, theory is the manner in which newly discovered and unanticipated facts arrange themselves under it. No more striking illustration of this can be found than in the contrasted fates of Cuvier's theory and of that of Darwin. Even before Cuvier's death his views had been undermined and the progress of discovery soon laid them in irreparable ruin, while the activity of half-a-century in many different lines of inquiry has established the theory of evolution upon a foundation of ever growing solidity. It is Darwin's imperishable glory that he prescribed the lines along which all the biological sciences were to advance to conquests not dreamed of when he wrote.



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