GEOLOGICAL CONTEMPORANEITY AND PERSISTENT TYPES OF LIFE.

By Thomas H. Huxley

1

MERCHANTS occasionally go through a wholesome, though troublesome and not always satisfactory, process which they term "taking stock." After all the excitement of speculation, the pleasure of gain, and the pain of loss, the trader makes up his mind to face facts and to learn the exact quantity and quality of his solid and reliable possessions.

The man of science does well sometimes to imitate this procedure; and, forgetting for the time the importance of his own small winnings, to re-examine the common stock in trade, so that he may make sure how far the stock of bullion in the cellar—on the faith of whose existence so much paper has been circulating—is really the solid gold of truth.

The Anniversary Meeting of the Geological Society seems to be an occasion well suited for an undertaking of this kind—for an inquiry, in fact, into the nature and value of the present results of paleontological investigation; and the more so, as all those who have paid close attention to the late multitudinous discussions in which paleontology is implicated, must have felt the urgent necessity of some such scrutiny.

First in order, as the most definite and unquestionable of all the results of paleontology, must be mentioned the immense extension and impulse given to botany, zoology, and comparative anatomy, by the investigation of fossil remains. Indeed, the mass of biological facts has been so greatly increased, and the range of biological speculation has been so vastly widened, by the researches of the geologist and paleontologist, that it is to be feared there are naturalists in existence who look upon geology as Brindley regarded rivers. "Rivers," said the great engineer, "were made to feed canals"; and geology, some seem to think, was solely created to advance comparative anatomy.

Were such a thought justifiable, it could hardly expect to be received with favour by this assembly. But it is not justifiable. Your favourite science has her own great aims independent of all others; and if, notwithstanding her steady devotion to her own progress, she can scatter such rich alms among her sisters, it should be remembered that her charity is of the sort that does not impoverish, but "blesseth him that gives and him that takes."

Regard the matter as we will, however, the facts remain. Nearly 40,000 species of animals and plants have been added to the Systema Naturae by paleontologic research. This is a living population equivalent to that of a new continent in mere number; equivalent to that of a new hemisphere, if we take into account the small population of insects as yet found fossil, and the large proportion and peculiar organization of many of the Vertebrata.

But, beyond this, it is perhaps not too much to say that, except for the necessity of interpreting paleontologic facts, the laws of distribution would have received less careful study; while few comparative anatomists (and those not of the first order) would have been induced by mere love of detail, as such, to study the minutiae of osteology, were it not that in such minutiae lie the only keys to the most interesting riddles offered by the extinct animal world.

These assuredly are great and solid gains. Surely it is matter for no small congratulation that in half a century (for paleontology, though it dawned earlier, came into full day only with Cuvier) a subordinate branch of biology should have doubled the value and the interest of the whole group of sciences to which it belongs.

But this is not all. Allied with geology, paleontology has established two laws of inestimable importance: the first, that one and the same area of the earth's surface has been successively occupied by very different kinds of living beings; the second, that the order of succession established in one locality holds good, approximately, in all.

The first of these laws is universal and irreversible; the second is an induction from a vast number of observations, though it may possibly, and even probably, have to admit of exceptions. As a consequence of the second law, it follows that a peculiar relation frequently subsists between series of strata, containing organic remains, in different localities. The series resemble one another, not only in virtue of a general resemblance of the organic remains in the two, but also in virtue of a resemblance in the order and character of the serial succession in each. There is a resemblance of arrangement; so that the separate terms of each series, as well as the whole series, exhibit a correspondence.

Succession implies time; the lower members of a series of sedimentary rocks are certainly older than the upper; and when the notion of age was once introduced as the equivalent of succession, it was no wonder that correspondence in succession came to be looked upon as a correspondence in age, or "contemporaneity." And, indeed, so long as relative age only is spoken of, correspondence in succession 'is' correspondence in age; it is 'relative' contemporaneity.

But it would have been very much better for geology if so loose and ambiguous a word as "contemporaneous" had been excluded from her terminology, and if, in its stead, some term expressing similarity of serial relation, and excluding the notion of time altogether, had been employed to denote correspondence in position in two or more series of strata.

In anatomy, where such correspondence of position has constantly to be spoken of, it is denoted by the word "homology" and its derivatives; and for Geology (which after all is only the anatomy and physiology of the earth) it might be well to invent some single word, such as "homotaxis" (similarity of order), in order to express an essentially similar idea. This, however, has not been done, and most probably the inquiry will at once be made—To what end burden science with a new and strange term in place of one old, familiar, and part of our common language?

The reply to this question will become obvious as the inquiry into the results of paleontology is pushed further.

Those whose business it is to acquaint themselves specially with the works of paleontologists, in fact, will be fully aware that very few, if any, would rest satisfied with such a statement of the conclusions of their branch of biology as that which has just been given.

Our standard repertories of paleontology profess to teach us far higher things—to disclose the entire succession of living forms upon the surface of the globe; to tell us of a wholly different distribution of climatic conditions in ancient times; to reveal the character of the first of all living existences; and to trace out the law of progress from them to us.

It may not be unprofitable to bestow on these professions a somewhat more critical examination than they have hitherto received, in order to ascertain how far they rest on an irrefragable basis; or whether, after all, it might not be well for paleontologists to learn a little more carefully that scientific "ars artium," the art of saying "I don't know." And to this end let us define somewhat more exactly the extent of these pretensions of paleontology.

Every one is aware that Professor Bronn's 'Untersuchungen' and Professor Pictet's 'Traite de Paleontologie' are works of standard authority, familiarly consulted by every working paleontologist. It is desirable to speak of these excellent books, and of their distinguished authors, with the utmost respect, and in a tone as far as possible removed from carping criticism; indeed, if they are specially cited in this place, it is merely in justification of the assertion that the following propositions, which may be found implicitly, or explicitly, in the works in question, are regarded by the mass of paleontologists and geologists, not only on the Continent but in this country, as expressing some of the best-established results of paleontology. Thus:—

Animals and plants began their existence together, not long after the commencement of the deposition of the sedimentary rocks; and then succeeded one another, in such a manner, that totally distinct faunae and florae occupied the whole surface of the earth, one after the other, and during distinct epochs of time.

A geological formation is the sum of all the strata deposited over the whole surface of the earth during one of these epochs: a geological fauna or flora is the sum of all the species of animals or plants which occupied the whole surface of the globe, during one of these epochs.

The population of the earth's surface was at first very similar in all parts, and only from the middle of the Tertiary epoch onwards, began to show a distinct distribution in zones.

The constitution of the original population, as well as the numerical proportions of its members, indicates a warmer and, on the whole, somewhat tropical climate, which remained tolerably equable throughout the year. The subsequent distribution of living beings in zones is the result of a gradual lowering of the general temperature, which first began to be felt at the poles.

It is not now proposed to inquire whether these doctrines are true or false; but to direct your attention to a much simpler though very essential preliminary question—What is their logical basis? what are the fundamental assumptions upon which they all logically depend? and what is the evidence on which those fundamental propositions demand our assent?

These assumptions are two: the first, that the commencement of the geological record is coeval with the commencement of life on the globe; the second, that geological contemporaneity is the same thing as chronological synchrony. Without the first of these assumptions there would of course be no ground for any statement respecting the commencement of life; without the second, all the other statements cited, every one of which implies a knowledge of the state of different parts of the earth at one and the same time, will be no less devoid of demonstration.

The first assumption obviously rests entirely on negative evidence. This is, of course, the only evidence that ever can be available to prove the commencement of any series of phenomena; but, at the same time, it must be recollected that the value of negative evidence depends entirely on the amount of positive corroboration it receives. If A B wishes to prove an 'alibi', it is of no use for him to get a thousand witnesses simply to swear that they did not see him in such and such a place, unless the witnesses are prepared to prove that they must have seen him had he been there. But the evidence that animal life commenced with the Lingula-flags, 'e.g.', would seem to be exactly of this unsatisfactory uncorroborated sort. The Cambrian witnesses simply swear they "haven't seen anybody their way"; upon which the counsel for the other side immediately puts in ten or twelve thousand feet of Devonian sandstones to make oath they never saw a fish or a mollusk, though all the world knows there were plenty in their time.

But then it is urged that, though the Devonian rocks in one part of the world exhibit no fossils, in another they do, while the lower Cambrian rocks nowhere exhibit fossils, and hence no living being could have existed in their epoch.

To this there are two replies: the first, that the observational basis of the assertion that the lowest rocks are nowhere fossiliferous is an amazingly small one, seeing how very small an area, in comparison to that of the whole world, has yet been fully searched; the second, that the argument is good for nothing unless the unfossiliferous rocks in question were not only 'contemporaneous' in the geological sense, but 'synchronous' in the chronological sense. To use the 'alibi' illustration again. If a man wishes to prove he was in neither of two places, A and B, on a given day, his witnesses for each place must be prepared to answer for the whole day. If they can only prove that he was not at A in the morning, and not at B in the afternoon, the evidence of his absence from both is 'nil', because he might have been at B in the morning and at A in the afternoon.

Thus everything depends upon the validity of the second assumption. And we must proceed to inquire what is the real meaning of the word "contemporaneous" as employed by geologists. To this end a concrete example may be taken.

The Lias of England and the Lias of Germany, the Cretaceous rocks of Britain and the Cretaceous rocks of Southern India, are termed by geologists "contemporaneous" formations; but whenever any thoughtful geologist is asked whether he means to say that they were deposited synchronously, he says, "No,—only within the same great epoch." And if, in pursuing the inquiry, he is asked what may be the approximate value in time of a "great epoch"—whether it means a hundred years, or a thousand, or a million, or ten million years—his reply is, "I cannot tell."

If the further question be put, whether physical geology is in possession of any method by which the actual synchrony (or the reverse) of any two distant deposits can be ascertained, no such method can be heard of; it being admitted by all the best authorities that neither similarity of mineral composition, nor of physical character, nor even direct continuity of stratum, are 'absolute' proofs of the synchronism of even approximated sedimentary strata: while, for distant deposits, there seems to be no kind of physical evidence attainable of a nature competent to decide whether such deposits were formed simultaneously, or whether they possess any given difference of antiquity. To return to an example already given: All competent authorities will probably assent to the proposition that physical geology does not enable us in any way to reply to this question—Were the British Cretaceous rocks deposited at the same time as those of India, or are they a million of years younger or a million of years older?

Is paleontology able to succeed where physical geology fails? Standard writers on paleontology, as has been seen, assume that she can. They take it for granted, that deposits containing similar organic remains are synchronous—at any rate in a broad sense; and yet, those who will study the eleventh and twelfth chapters of Sir Henry De La Beche's remarkable 'Researches in Theoretical Geology', published now nearly thirty years ago, and will carry out the arguments there most luminously stated, to their logical consequences, may very easily convince themselves that even absolute identity of organic contents is no proof of the synchrony of deposits, while absolute diversity is no proof of difference of date. Sir Henry De La Beche goes even further, and adduces conclusive evidence to show that the different parts of one and the same stratum, having a similar composition throughout, containing the same organic remains, and having similar beds above and below it, may yet differ to any conceivable extent in age.

Edward Forbes was in the habit of asserting that the similarity of the organic contents of distant formations was 'prima facie' evidence, not of their similarity, but of their difference of age; and holding as he did the doctrine of single specific centres, the conclusion was as legitimate as any other; for the two districts must have been occupied by migration from one of the two, or from an intermediate spot, and the chances against exact coincidence of migration and of imbedding are infinite.

In point of fact, however, whether the hypothesis of single or of multiple specific centres be adopted, similarity of organic contents cannot possibly afford any proof of the synchrony of the deposits which contain them; on the contrary, it is demonstrably compatible with the lapse of the most prodigious intervals of time, and with the interposition of vast changes in the organic and inorganic worlds, between the epochs in which such deposits were formed.

On what amount of similarity of their faunae is the doctrine of the contemporaneity of the European and of the North American Silurians based? In the last edition of Sir Charles Lyell's 'Elementary Geology' it is stated, on the authority of a former President of this Society, the late Daniel Sharpe, that between 30 and 40 per cent. of the species of Silurian Mollusca are common to both sides of the Atlantic. By way of due allowance for further discovery, let us double the lesser number and suppose that 60 per cent. of the species are common to the North American and the British Silurians. Sixty per cent. of species in common is, then, proof of contemporaneity.

Now suppose that, a million or two of years hence, when Britain has made another dip beneath the sea and has come up again, some geologist applies this doctrine, in comparing the strata laid bare by the upheaval of the bottom, say, of St. George's Channel with what may then remain of the Suffolk Crag. Reasoning in the same way, he will at once decide the Suffolk Crag and the St. George's Channel beds to be contemporaneous; although we happen to know that a vast period (even in the geological sense) of time, and physical changes of almost unprecedented extent, separate the two.

But if it be a demonstrable fact that strata containing more than 60 or 70 per cent. of species of Mollusca in common, and comparatively close together, may yet be separated by an amount of geological time sufficient to allow of some of the greatest physical changes the world has seen, what becomes of that sort of contemporaneity the sole evidence of which is a similarity of facies, or the identity of half a dozen species, or of a good many genera?

And yet there is no better evidence for the contemporaneity assumed by all who adopt the hypothesis of universal faunae and florae, of a universally uniform climate, and of a sensible cooling of the globe during geological time.

There seems, then, no escape from the admission that neither physical geology, nor paleontology, possesses any method by which the absolute synchronism of two strata can be demonstrated. All that geology can prove is local order of succession. It is mathematically certain that, in any given vertical linear section of an undisturbed series of sedimentary deposits, the bed which lies lowest is the oldest. In many other vertical linear sections of the same series, of course, corresponding beds will occur in a similar order; but, however great may be the probability, no man can say with absolute certainty that the beds in the two sections were synchronously deposited. For areas of moderate extent, it is doubtless true that no practical evil is likely to result from assuming the corresponding beds to be synchronous or strictly contemporaneous; and there are multitudes of accessory circumstances which may fully justify the assumption of such synchrony. But the moment the geologist has to deal with large areas, or with completely separated deposits, the mischief of confounding that "homotaxis" or "similarity of arrangement," which 'can' be demonstrated, with "synchrony" or "identity of date," for which there is not a shadow of proof, under the one common term of "contemporaneity" becomes incalculable, and proves the constant source of gratuitous speculations.

For anything that geology or paleontology are able to show to the contrary, a Devonian fauna and flora in the British Islands may have been contemporaneous with Silurian life in North America, and with a Carboniferous fauna and flora in Africa. Geographical provinces and zones may have been as distinctly marked in the Paleozoic epoch as at present, and those seemingly sudden appearances of new genera and species, which we ascribe to new creation, may be simple results of migration.

It may be so; it may be otherwise. In the present condition of our knowledge and of our methods, one verdict—"not proven, and not provable"—must be recorded against all the grand hypotheses of the paleontologist respecting the general succession of life on the globe. The order and nature of terrestrial life, as a whole, are open questions. Geology at present provides us with most valuable topographical records, but she has not the means of working them into a universal history. Is such a universal history, then, to be regarded as unattainable? Are all the grandest and most interesting problems which offer themselves to the geological student essentially insoluble? Is he in the position of a scientific Tantalus—doomed always to thirst for a knowledge which he cannot obtain? The reverse is to be hoped; nay, it may not be impossible to indicate the source whence help will come.

In commencing these remarks, mention was made of the great obligations under which the naturalist lies to the geologist and paleontologist. Assuredly the time will come when these obligations will be repaid tenfold, and when the maze of the world's past history, through which the pure geologist and the pure paleontologist find no guidance, will be securely threaded by the clue furnished by the naturalist.

All who are competent to express an opinion on the subject are, at present, agreed that the manifold varieties of animal and vegetable form have not either come into existence by chance, nor result from capricious exertions of creative power; but that they have taken place in a definite order, the statement of which order is what men of science term a natural law. Whether such a law is to be regarded as an expression of the mode of operation of natural forces, or whether it is simply a statement of the manner in which a supernatural power has thought fit to act, is a secondary question, so long as the existence of the law and the possibility of its discovery by the human intellect are granted. But he must be a half-hearted philosopher who, believing in that possibility, and having watched the gigantic strides of the biological sciences during the last twenty years, doubts that science will sooner or later make this further step, so as to become possessed of the law of evolution of organic forms—of the unvarying order of that great chain of causes and effects of which all organic forms, ancient and modern, are the links. And then, if ever, we shall be able to begin to discuss, with profit, the questions respecting the commencement of life, and the nature of the successive populations of the globe, which so many seem to think are already answered.

The preceding arguments make no particular claim to novelty; indeed they have been floating more or less distinctly before the minds of geologists for the last thirty years; and if, at the present time, it has seemed desirable to give them more definite and systematic expression, it is because paleontology is every day assuming a greater importance, and now requires to rest on a basis the firmness of which is thoroughly well assured. Among its fundamental conceptions, there must be no confusion between what is certain and what is more or less probable. 2 But, pending the construction of a surer foundation than paleontology now possesses, it may be instructive, assuming for the nonce the general correctness of the ordinary hypothesis of geological contemporaneity, to consider whether the deductions which are ordinarily drawn from the whole body of paleontologic facts are justifiable.

The evidence on which such conclusions are based is of two kinds, negative and positive. The value of negative evidence, in connection with this inquiry, has been so fully and clearly discussed in an address from the chair of this Society 3, which none of us have forgotten, that nothing need at present be said about it; the more, as the considerations which have been laid before you have certainly not tended to increase your estimation of such evidence. It will be preferable to turn to the positive facts of paleontology, and to inquire what they tell us.

We are all accustomed to speak of the number and the extent of the changes in the living population of the globe during geological time as something enormous: and indeed they are so, if we regard only the negative differences which separate the older rocks from the more modern, and if we look upon specific and generic changes as great changes, which from one point of view, they truly are. But leaving the negative differences out of consideration, and looking only at the positive data furnished by the fossil world from a broader point of view—from that of the comparative anatomist who has made the study of the greater modifications of animal form his chief business—a surprise of another kind dawns upon the mind; and under 'this' aspect the smallness of the total change becomes as astonishing as was its greatness under the other.

There are two hundred known orders of plants; of these not one is certainly known to exist exclusively in the fossil state. The whole lapse of geological time has as yet yielded not a single new ordinal type of vegetable structure. 4

The positive change in passing from the recent to the ancient animal world is greater, but still singularly small. No fossil animal is so distinct from those now living as to require to be arranged even in a separate class from those which contain existing forms. It is only when we come to the orders, which may be roughly estimated at about a hundred and thirty, that we meet with fossil animals so distinct from those now living as to require orders for themselves; and these do not amount, on the most liberal estimate, to more than about 10 per cent. of the whole.

There is no certainly known extinct order of Protozoa; there is but one among the Coelenterata—that of the rugose corals; there is none among the Mollusca; there are three, the Cystidea, Blastoidea, and Edrioasterida, among the Echinoderms; and two, the Trilobita and Eurypterida, among the Crustacea; making altogether five for the great sub-kingdom of Annulosa. Among Vertebrates there is no ordinally distinct fossil fish: there is only one extinct order of Amphibia—the Labyrinthodonts; but there are at least four distinct orders of Reptilia, viz. the Ichthyosauria, Plesiosauria, Pterosauria, Dinosauria, and perhaps another or two. There is no known extinct order of Birds, and no certainly known extinct order of Mammals, the ordinal distinctness of the "Toxodontia" being doubtful.

The objection that broad statements of this kind, after all, rest largely on negative evidence is obvious, but it has less force than may at first be supposed; for, as might be expected from the circumstances of the case, we possess more abundant positive evidence regarding Fishes and marine Mollusks than respecting any other forms of animal life; and yet these offer us, through the whole range of geological time, no species ordinally distinct from those now living; while the far less numerous class of Echinoderms presents three; and the Crustacea two, such orders, though none of these come down later than the Paleozoic age. Lastly, the Reptilia present the extraordinary and exceptional phenomenon of as many extinct as existing orders, if not more; the four mentioned maintaining their existence from the Lias to the Chalk inclusive.

Some years ago one of your Secretaries pointed out another kind of positive paleontologic evidence tending towards the same conclusion—afforded by the existence of what he termed "persistent types" of vegetable and of animal life. 5 He stated, on the authority of Dr. Hooker, that there are Carboniferous plants which appear to be generically identical with some now living; that the cone of the Oolitic 'Araucaria' is hardly distinguishable from that of an existing species; that a true 'Pinus' appears in the Purbecks, and a 'Juglans' in the Chalk; while, from the Bagshot Sands, a 'Banksia', the wood of which is not distinguishable from that of species now living in Australia, had been obtained.

Turning to the animal kingdom, he affirmed the tabulate corals of the Silurian rocks to be wonderfully like those which now exist; while even the families of the Aporosa were all represented in the older Mesozoic rocks.

Among the Molluska similar facts were adduced. Let it be borne in mind that 'Avicula', 'Mytails', 'Chiton', 'Natica', 'Patella', 'Trochus', 'Discina', 'Orbicula', 'Lingula', 'Rhynchonella', and 'Nautilus', all of which are existing 'genera', are given without a doubt as Silurian in the last edition of 'Siluria'; while the highest forms of the highest Cephalopods are represented in the Lias by a genus, 'Belemnoteuthis', which presents the closest relation to the existing 'Loligo'.

The two highest groups of the Annulosa, the Insecta and the Arachnida, are represented in the Coal, either by existing genera, or by forms differing from existing genera in quite minor peculiarities.

Turning to the Vertebrata, the only Paleozoic Elasmobranch Fish of which we have any complete knowledge is the Devonian and Carboniferous 'Pleuracanthus', which differs no more from existing Sharks than these do from one another.

Again, vast as is the number of undoubtedly Ganoid fossil Fishes, and great as is their range in time, a large mass of evidence has recently been adduced to show that almost all those respecting which we possess sufficient information, are referable to the same sub-ordinal groups as the existing 'Lepidosteus', 'Polypterus', and Sturgeon; and that a singular relation obtains between the older and the younger Fishes; the former, the Devonian Ganoids, being almost all members of the same sub-order as 'Polypterus', while the Mesozoic Ganoids are almost all similarly allied to 'Lepidosteus'. 6

Again, what can be more remarkable than the singular constancy of structure preserved throughout a vast period of time by the family of the Pycnodonts and by that of the true Coelacanths; the former persisting, with but insignificant modifications, from the Carboniferous to the Tertiary rocks, inclusive; the latter existing, with still less change, from the Carboniferous rocks to the Chalk, inclusive?

Among Reptiles, the highest living group, that of the Crocodilia, is represented, at the early part of the Mesozoic epoch, by species identical in the essential characters of their organization with those now living, and differing from the latter only in such matters as the form of the articular facets of the vertebral centra, in the extent to which the nasal passages are separated from the cavity of the mouth by bone, and in the proportions of the limbs.

And even as regards the Mammalia, the scanty remains of Triassic and Oolitic species afford no foundation for the supposition that the organization of the oldest forms differed nearly so much from some of those which now live as these differ from one another.

It is needless to multiply these instances; enough has been said to justify the statement that, in view of the immense diversity of known animal and vegetable forms, and the enormous lapse of time indicated by the accumulation of fossiliferous strata, the only circumstance to be wondered at is, not that the changes of life, as exhibited by positive evidence, have been so great, but that they have been so small.

Be they great or small, however, it is desirable to attempt to estimate them. Let us, therefore, take each great division of the animal world in succession, and, whenever an order or a family can be shown to have had a prolonged existence, let us endeavour to ascertain how far the later members of the group differ from the earlier ones. If these later members, in all or in many cases, exhibit a certain amount of modification, the fact is, so far, evidence in favour of a general law of change; and, in a rough way, the rapidity of that change will be measured by the demonstrable amount of modification. On the other hand, it must be recollected that the absence of any modification, while it may leave the doctrine of the existence of a law of change without positive support, cannot possibly disprove all forms of that doctrine, though it may afford a sufficient refutation of any of them.

The PROTOZOA.—The Protozoa are represented throughout the whole range of geological series, from the Lower Silurian formation to the present day. The most ancient forms recently made known by Ehrenberg are exceedingly like those which now exist: no one has ever pretended that the difference between any ancient and any modern Foraminifera is of more than generic value, nor are the oldest Foraminifera either simpler, more embryonic, or less differentiated, than the existing forms.

The COELENTERATA.—The Tabulate Corals have existed from the Silurian epoch to the present day, but I am not aware that the ancient 'Heliolites' possesses a single mark of a more embryonic or less differentiated character, or less high organization, than the existing 'Heliopora'. As for the Aporose Corals, in what respect is the Silurian 'Paleocyclus' less highly organized or more embryonic than the modern 'Fungia', or the Liassic Aporosa than the existing members of the same families?

The 'Mollusca'.—In what sense is the living 'Waldheimia' less embryonic, or more specialized; than the paleozoic 'Spirifer'; or the existing 'Rhynchonellae', 'Craniae', 'Discinae', 'Lingulae', than the Silurian species of the same genera? In what sense can 'Loligo' or 'Spirula' be said to be more specialized, or less embryonic, than 'Belemnites'; or the modern species of Lamellibranch and Gasteropod genera, than the Silurian species of the same genera?

The ANNULOSA.—The Carboniferous Insecta and Arachnida are neither less specialized, nor more embryonic, than these that now live, nor are the Liassic Cirripedia and Macrura; while several of the Brachyura, which appear in the Chalk, belong to existing genera; and none exhibit either an intermediate, or an embryonic, character.

The VERTEBRARA.—Among fishes I have referred to the Coelacanthini (comprising the genera 'Coelacanthus', 'Holophagus', 'Undina', and 'Macropoma') as affording an example of a persistent type; and it is most remarkable to note the smallness of the differences between any of these fishes (affecting at most the proportions of the body and fins, and the character and sculpture of the scales), notwithstanding their enormous range in time. In all the essentials of its very peculiar structure, the 'Macropoma' of the Chalk is identical with the 'Coelacanthus' of the Coal. Look at the genus 'Lepidotus', again, persisting without a modification of importance from the Liassic to the Eocene formations inclusive.

Or among the Teleostei—in what respect is the 'Beryx' of the Chalk more embryonic, or less differentiated, than 'Beryx lineatus' of King George's Sound?

Or to turn to the higher Vertebrata—in what sense are the Liassic Chelonia inferior to those which now exist? How are the Cretaceous Ichthyosauria, Plesiosauria, or Pterosauria less embryonic, or more differentiated, species than those of the Lias?

Or lastly, in what circumstance is the 'Phascolotherium' more embryonic, or of a more generalized type, than the modern Opossum; or a 'Lophiodon', or a 'Paleotherium', than a modern 'Tapirus' or 'Hyrax'?

These examples might be almost indefinitely multiplied, but surely they are sufficient to prove that the only safe and unquestionable testimony we can procure—positive evidence—fails to demonstrate any sort of progressive modification towards a less embryonic, or less generalised, type in a great many groups of animals of long-continued geological existence. In these groups there is abundant evidence of variation—none of what is ordinarily understood as progression; and, if the known geological record is to be regarded as even any considerable fragment of the whole, it is inconceivable that any theory of a necessarily progressive development can stand, for the numerous orders and families cited afford no trace of such a process.

But it is a most remarkable fact, that, while the groups which have been mentioned, and many besides, exhibit no sign of progressive modification, there are others, co-existing with them, under the same conditions, in which more or less distinct indications of such a process seems to be traceable. Among such indications I may remind you of the predominance of Holostome Gasteropoda in the older rocks as compared with that of Siphonostome Gasteropoda in the later. A case less open to the objection of negative evidence, however, is that afforded by the Tetrabranchiate Cephalopoda, the forms of the shells and of the septal sutures exhibiting a certain increase of complexity in the newer genera. Here, however, one is met at once with the occurrence of 'Orthoceras' and 'Baculites' at the two ends of the series, and of the fact that one of the simplest Genera, 'Nautilus', is that which now exists.

The Crinoidea, in the abundance of stalked forms in the ancient formations as compared with their present rarity, seem to present us with a fair case of modification from a more embryonic towards a less embryonic condition. But then, on careful consideration of the facts, the objection arises that the stalk, calyx, and arms of the paleozoic Crinoid are exceedingly different from the corresponding organs of a larval 'Comatula'; and it might with perfect justice be argued that 'Actinocrinus' and 'Eucalyptocrinus', for example, depart to the full as widely, in one direction, from the stalked embryo of 'Comatula', as 'Comatula' itself does in the other.

The Echinidea, again, are frequently quoted as exhibiting a gradual passage from a more generalized to a more specialized type, seeing that the elongated, or oval, Spatangoids appear after the spheroidal Echinoids. But here it might be argued, on the other hand, that the spheroidal Echinoids, in reality, depart further from the general plan and from the embryonic form than the elongated Spatangoids do; and that the peculiar dental apparatus and the pedicellariae of the former are marks of at least as great differentiation as the petaloid ambulacra and semitae of the latter.

Once more, the prevalence of Macrurous before Brachyurous Podophthalmia is, apparently, a fair piece of evidence in favour of progressive modification in the same order of Crustacea; and yet the case will not stand much sifting, seeing that the Macrurous Podophthalmia depart as far in one direction from the common type of Podophthalmia, or from any embryonic condition of the Brachyura, as the Brachyura do in the other; and that the middle terms between Macrura and Brachyura—the Anomura—are little better represented in the older Mesozoic rocks than the Brachyura are.

None of the cases of progressive modification which are cited from among the Invertebrata appear to me to have a foundation less open to criticism than these; and if this be so, no careful reasoner would, I think, be inclined to lay very great stress upon them. Among the Vertebrata, however, there are a few examples which appear to be far less open to objection.

It is, in fact, true of several groups of Vertebrata which have lived through a considerable range of time, that the endoskeleton (more particularly the spinal column) of the older genera presents a less ossified, and, so far, less differentiated, condition than that of the younger genera. Thus the Devonian Ganoids, though almost all members of the same sub-order as 'Polypterus', and presenting numerous important resemblances to the existing genus, which possesses biconcave vertebrae, are, for the most part, wholly devoid of ossified vertebral centra. The Mesozoic Lepidosteidae, again, have, at most, biconcave vertebrae, while the existing 'Lepidosteus' has Salamandroid, opisthocoelous, vertebrae. So, none of the Paleozoic Sharks have shown themselves to be possessed of ossified vertebrae, while the majority of modern Sharks possess such vertebrae. Again, the more ancient Crocodilia and Lacertilia have vertebrae with the articular facets of their centra flattened or biconcave, while the modern members of the same group have them procoelous. But the most remarkable examples of progressive modification of the vertebral column, in correspondence with geological age, are those afforded by the Pycnodonts among fish, and the Labyrinthodonts among Amphibia.

The late able ichthyologist Heckel pointed out the fact, that, while the Pycnodonts never possess true vertebral centra, they differ in the degree of expansion and extension of the ends of the bony arches of the vertebrae upon the sheath of the notochord; the Carboniferous forms exhibiting hardly any such expansion, while the Mesozoic genera present a greater and greater development, until, in the Tertiary forms, the expanded ends become suturally united so as to form a sort of false vertebra. Hermann von Meyer, again, to whose luminous researches we are indebted for our present large knowledge of the organization of the older Labyrinthodonts, has proved that the Carboniferous 'Archegosaurus' had very imperfectly developed vertebral centra, while the Triassic 'Mastodonsaurus' had the same parts completely ossified. 7

The regularity and evenness of the dentition of the 'Anoplotherium', as contrasted with that of existing Artiodactyles, and the assumed nearer approach of the dentition of certain ancient Carnivores to the typical arrangement, have also been cited as exemplifications of a law of progressive development, but I know of no other cases based on positive evidence which are worthy of particular notice.

What, then, does an impartial survey of the positively ascertained truths of paleontology testify in relation to the common doctrines of progressive modification, which suppose that modification to have taken place by a necessary progress from more to less embryonic forms, or from more to less generalized types, within the limits of the period represented by the fossiliferous rocks?

It negatives those doctrines; for it either shows us no evidence of any such modification, or demonstrates it to have been very slight; and as to the nature of that modification, it yields no evidence whatsoever that the earlier members of any long-continued group were more generalized in structure than the later ones. To a certain extent, indeed, it may be said that imperfect ossification of the vertebral column is an embryonic character; but, on the other hand, it would be extremely incorrect to suppose that the vertebral columns of the older Vertebrata are in any sense embryonic in their whole structure.

Obviously, if the earliest fossiliferous rocks now known are coeval with the commencement of life, and if their contents give us any just conception of the nature and the extent of the earliest fauna and flora, the insignificant amount of modification which can be demonstrated to have taken place in any one group of animals, or plants, is quite incompatible with the hypothesis that all living forms are the results of a necessary process of progressive development, entirely comprised within the time represented by the fossiliferous rocks.

Contrariwise, any admissible hypothesis of progressive modification must be compatible with persistence without progression, through indefinite periods. And should such an hypothesis eventually be proved to be true, in the only way in which it can be demonstrated, viz. by observation and experiment upon the existing forms of life, the conclusion will inevitably present itself, that the Paleozoic, Mesozoic, and Cainozoic faunae and florae, taken together, bear somewhat the same proportion to the whole series of living beings which have occupied this globe, as the existing fauna and flora do to them.

Such are the results of paleontology as they appear, and have for some years appeared, to the mind of an inquirer who regards that study simply as one of the applications of the great biological sciences, and who desires to see it placed upon the same sound basis as other branches of physical inquiry. If the arguments which have been brought forward are valid, probably no one, in view of the present state of opinion, will be inclined to think the time wasted which has been spent upon their elaboration.

1 (return)
[ The Anniversary Address to the Geological Society for 1862.]

2 (return)
[ "le plus grand service qu'on puisse rendre a la science est d'y faire place nette avant d'y rien construire."—CUVIER]

3 (return)
[ Anniversary Address for 1851, 'Quart. Journ. Geol. Soc.' vol. vii.]

4 (return)
[ See Hooker's 'Introductory Essay to the Flora of Tasmania', p. xxiii.]

5 (return)
[ See the abstract of a Lecture "On the Persistent Types of Animal Life," in the 'Notices of the Meetings of the Royal Institution of Great Britain'.—June 3, 1859, vol. iii. p. 151.

6 (return)
[ "Memoirs of the Geological Survey of the United Kingdom.—Decade x. Preliminary Essay upon the Systematic Arrangement of the Fishes of the Devonian Epoch."]

7 (return)
[ As the Address is passing through the press (March 7, 1862), evidence lies before me of the existence of a new Labyrinthodont ('Pholidogaster'), from the Edinburgh coal-field, with well-ossified vertebral centra.]