Discourses: Biological and Geological — Palaentology and the Doctrine of Evolution Pt 4

Let us now take a step further back in time, and inquire into the relations between the Miocene Fauna and its predecessor of the Upper Eocene formation.

Here it is to be regretted that our materials for forming a judgment are nothing to be compared in point of extent or variety with those which are yielded by the Miocene strata.

However, what we do know of this Upper Eocene Fauna of Europe gives sufficient positive information to enable us to draw some tolerably safe inferences.

It has yielded representatives of _Insectivora_, of _Cheiroptera_, of _Rodentia_, of _Carnivora_, of artiodactyle and perissodactyle _Ungulata_, and of opossum-like Marsupials.

No Australian type of Marsupial has been discovered in the Upper Eocene strata, nor any Edentate mammal.

The genera (except perhaps in the case of some of the _Insectivora, Cheiroptera_, and _Rodentia_) are different from those of the Miocene epoch, but present a remarkable general similarity to the Miocene and recent genera.

In several cases, as I have already shown, it has now been clearly made out that the relation between the Eocene and Miocene forms is such that the Eocene form is the less specialised; while its Miocene ally is more so, and the specialisation reaches its maximum in the recent forms of the same type.

So far as the Upper Eocene and the Miocene Mammalian Faunae are comparable, their relations are such as in no way to oppose the hypothesis that the older are the progenitors of the more recent forms, while, in some cases, they distinctly favour that hypothesis.

The period in tine and the changes in physical geography represented by the nummulitic deposits are undoubtedly very great, while the remains of Middle Eocene and Older Eocene Mammals are comparatively few.

The general facies of the Middle Eocene Fauna, however, is quite that of the Upper.

The Older Eocene pre-nummulitic mammalian Fauna contains Bats, two genera of _Carivora_, three genera of _Ungulata_ (probably all perissodactyle), and a didelphid Marsupial; all these forms, except perhaps the Bat and the Opossum, belong to genera which are not known to occur out of the Lower Eocene formation.

The _Coryphodon_ appears to have been allied to the Miocene and later Tapirs, while _Pliolophus_, in its skull and dentition, curiously partakes of both artiodactyle and perissodactyle characters; the third trochanter upon its femur, and its three-toed hind foot, however, appear definitely to fix its position in the latter division.

There is nothing, then, in what is known of the older Eocene mammals of the Arctogaeal province to forbid the supposition that they stood in an ancestral relation to those of the Calcaire Grossier and the Gypsum of the Paris basin, and that our present fauna, therefore, is directly derived from that which already existed in Arctogaea at the commencement of the Tertiary period.

But if we now cross the frontier between the Cainozoic and the Mesozoic faunae, as they are preserved within the Arctogaeal area, we meet with an astounding change, and what appears to be a complete and unmistakable break in the line of biological continuity.

Among the twelve or fourteen species of _Mammalia_ which are said to have been found in the Purbecks, not one is a member of the orders _Cheiroptera, Rodentia, Ungulata_, or _Carnivora_, which are so well represented in the Tertiaries.

No _Insectivora_ are certainly known, nor any opossum-like Marsupials.

Thus there is a vast negative difference between the Cainozoic and the Mesozoic mammalian faunae of Europe.

But there is a still more important positive difference, inasmuch as all these Mammalia appear to be Marsupials belonging to Australian groups, and thus appertaining to a different distributional province from the Eocene and Miocene marsupials, which are Austro-Columbian.

So far as the imperfect materials which exist enable a judgment to be formed, the same law appears to have held good for all the earlier Mesozoic _Mammalia_.

Of the Stonesfield slate mammals, one, _Amphitherium_, has a definitely Australian character; one, _Phascolotherium_, may be either Dasyurid or Didelphine; of a third, _Stereognathus_, nothing can at present be said.

The two mammals of the Trias, also, appear to belong to Australian groups.

Every one is aware of the many curious points of resemblance between the marine fauna of the European Mesozoic rocks and that which now exists in Australia.

But if there was this Australian facies about both the terrestrial and the marine faunae of Mesozoic Europe, and if there is this unaccountable and immense break between the fauna of Mesozoic and that of Tertiary Europe, is it not a very obvious suggestion that, in the Mesozoic epoch, the Australian province included Europe, and that the Arctogaeal province was contained within other limits?

The Arctogaeal province is at present enormous, while the Australian is relatively small.

Why should not these proportions have been different during the Mesozoic epoch?

Thus I am led to think that by far the simplest and most rational mode of accounting for the great change which took place in the living inhabitants of the European area at the end of the Mesozoic epoch, is the supposition that it arose from a vast alteration of the physical geography of the globe; whereby an area long tenanted by Cainozoic forms was brought into such relations with the European area that migration from the one to the other became possible, and took place on a great scale.

This supposition relieves us, at once, from the difficulty in which we were left, some time ago, by the arguments which I used to demonstrate the necessity of the existence of all the great types of the Eocene epoch in some antecedent period.

It is this Mesozoic continent (which may well have lain in the neighbourhood of what are now the shores of the North Pacific Ocean) which I suppose to have been occupied by the Mesozoic _Monodelphia_; and it is in this region that I conceive they must have gone through the long series of changes by which they were specialised into the forms which we refer to different orders.

I think it very probable that what is now South America may have received the characteristic elements of its mammalian fauna during the Mesozoic epoch; and there can be little doubt that the general nature of the change which took place at the end of the Mesozoic epoch in Europe was the upheaval of the eastern and northern regions of the Mesozoic sea-bottom into a westward extension of the Mesozoic continent, over which the mammalian fauna, by which it was already peopled, gradually spread.

This invasion of the land was prefaced by a previous invasion of the Cretaceous sea by modern forms of mollusca and fish.

It is easy to imagine how an analogous change might come about in the existing world.

There is, at present, a great difference between the fauna of the Polynesian Islands and that of the west coast of America.

The animals which are leaving their spoils in the deposits now forming in these localities are widely different.

Hence, if a gradual shifting of the deep sea, which at present bars migration between the easternmost of these islands and America, took place to the westward, while the American side of the sea-bottom was gradually upheaved, the palaeontologist of the future would find, over the Pacific area, exactly such a change as I am supposing to have occurred in the North-Atlantic area at the close of the Mesozoic period.

An Australian fauna would be found underlying an American fauna, and the transition from the one to the other would be as abrupt as that between the Chalk and lower Tertiaries; and as the drainage-area of the newly formed extension of the American continent gave rise to rivers and lakes, the mammals mired in their mud would differ from those of like deposits on the Australian side, just as the Eocene mammals differ from those of the Purbecks.

How do similar reasonings apply to the other great change of life--that which took place at the end of the Palaeozoic period?

In the Triassic epoch, the distribution of the dry land and of terrestrial vertebrate life appears to have been, generally, similar to that which existed in the Mesozoic epoch; so that the Triassic continents and their faunae seem to be related to the Mesozoic lands and their faunae, just as those of the Miocene epoch are related to those of the present day.

In fact, as I have recently endeavoured to prove to the Society, there was an Arctogaeal continent and an Arctogaeal province of distribution in Triassic times as there is now; and the _Sauropsida_ and _Marsupialia_ which constituted that fauna were, I doubt not, the progenitors of the _Sauropsida_ and _Marsupialia_ of the whole Mesozoic epoch.

Looking at the present terrestrial fauna of Australia, it appears to me to be very probable that it is essentially a remnant of the fauna of the Triassic, or even of an earlier, age[7] in which case Australia must at that time have been in continuity with the Arctogaeal continent.

[Footnote 7: Since this Address was read, Mr. Krefft has sent us news of the discovery in Australia of a freshwater fish of strangely Palaeozoic aspect, and apparently a Ganoid intermediate between _Dipterus_ and _Lepidosiren_.

[The now well-known _Ceratodus_.

1894. ]] But now comes the further inquiry, Where was the highly differentiated Sauropsidan fauna of the Trias in Palaeozoic times?

The supposition that the Dinosaurian, Crocodilian, Dicynodontian, and to Plesiosaurian types were suddenly created at the end of the Permian epoch may be dismissed, without further consideration, as a monstrous and unwarranted assumption.

The supposition that all these types were rapidly differentiated out of _Lacertilia_ in the time represented by the passage from the Palaeozoic to the Mesozoic formation, appears to me to be hardly more credible, to say nothing of the indications of the existence of Dinosaurian forms in the Permian rocks which have already been obtained.

For my part, I entertain no sort of doubt that the Reptiles, Birds, and Mammals of the Trias are the direct descendants of Reptiles, Birds, and Mammals which existed in the latter part of the Palaeozoic epoch, but not in any area of the present dry land which has yet been explored by the geologist.

This may seem a bold assumption, but it will not appear unwarrantable to those who reflect upon the very small extent of the earth's surface which has hitherto exhibited the remains of the great Mammalian fauna of the Eocene times.

In this respect, the Permian land Vertebrate fauna appears to me to be related to the Triassic much as the Eocene is to the Miocene.

Terrestrial reptiles have been found in Permian rocks only in three localities; in some spots of France, and recently of England, and over a more extensive area in Germany.

Who can suppose that the few fossils yet found in these regions give any sufficient representation of the Permian fauna?

It may be said that the Carboniferous formations demonstrate the existence of a vast extent of dry land in the present dry-land area, and that the supposed terrestrial Palaeozoic Vertebrate Fauna ought to have left its remains in the Coal-measures, especially as there is now reason to believe that much of the coal was formed by the accumulation of spores and sporangia on dry land.

But if we consider the matter more closely, I think that this apparent objection loses its force.

It is clear that, during the Carboniferous epoch, the vast area of land which is now covered by Coal-measures must have been undergoing a gradual depression.

The dry land thus depressed must, therefore, have existed, as such, before the Carboniferous epoch--in other words, in Devonian times--and its terrestrial population may never have been other than such as existed during the Devonian, or some previous epoch, although much higher forms may have been developed elsewhere.

Again, let me say that I am making no gratuitous assumption of inconceivable changes.

It is clear that the enormous area of Polynesia is, on the whole, an area over which depression has taken place to an immense extent; consequently a great continent, or assemblage of subcontinental masses of land must have existed at some former time, and that at a recent period, geologically speaking, in the area of the Pacific.

But if that continent had contained Mammals, some of them must have remained to tell the tale; and as it is well known that these islands have no indigenous _Mammalia_, it is safe to assume that none existed.

Thus, midway between Australia and South America, each of which possesses an abundant and diversified mammalian fauna, a mass of land, which may have been as large as both put together, must have existed without a mammalian inhabitant.

Suppose that the shores of this great land were fringed, as those of tropical Australia are now, with belts of mangroves, which would extend landwards on the one side, and be buried beneath littoral deposits on the other side, as depression went on; and great beds of mangrove lignite might accumulate over the sinking land.

Let upheaval of the whole now take place, in such a manner as to bring the emerging land into continuity with the South-American or Australian continent, and, in course of time, it would be peopled by an extension of the fauna of one of these two regions--just as I imagine the European Permian dry land to have been peopled.

I see nothing whatever against the supposition that distributional provinces of terrestrial life existed in the Devonian epoch, inasmuch as M.

Barrande has proved that they existed much earlier.

I am aware of no reason for doubting that, as regards the grades of terrestrial life contained in them, one of these may have been related to another as New Zealand is to Australia, or as Australia is to India, at the present day.

Analogy seems to me to be rather in favour of, than against, the supposition that while only Ganoid fishes inhabited the fresh waters of our Devonian land, _Amphibia_ and _Reptilia_, or even higher forms, may have existed, though we have not yet found them.

The earliest Carboniferous _Amphibia_ now known, such as _Anthracosaurus_, are so highly specialised that I can by no means conceive that they have been developed out of piscine forms in the interval between the Devonian and the Carboniferous periods, considerable as that is.

And I take refuge in one of two alternatives: either they existed in our own area during the Devonian epoch and we have simply not yet found them; or they formed part of the population of some other distributional province of that day, and only entered our area by migration at the end of the Devonian epoch.

Whether _Reptilia_ and _Mammalia_ existed along with them is to me, at present, a perfectly open question, which is just as likely to receive an affirmative as a negative answer from future inquirers.

Let me now gather together the threads of my argumentation into the form of a connected hypothetical view of the manner in which the distribution of living and extinct animals has been brought about.

I conceive that distinct provinces of the distribution of terrestrial life have existed since the earliest period at which that life is recorded, and possibly much earlier; and I suppose, with Mr. Darwin, that the progress of modification of terrestrial forms is more rapid in areas of elevation than in areas of depression.

I take it to be certain that Labyrinthodont _Amphibia_ existed in the distributional province which included the dry land depressed during the Carboniferous epoch; and I conceive that, in some other distributional provinces of that day, which remained in the condition of stationary or of increasing dry land, the various types of the terrestrial _Sauropsida_ and of the _Mammalia_ were gradually developing.

The Permian epoch marks the commencement of a new movement of upheaval in our area, which dry land existed in North America, Europe, Asia, and Africa, as it does now.

Into this great new continental area the Mammals, Birds, and Reptiles developed during the Palaeozoic epoch spread, and formed the great Triassic Arctogaeal province.

But, at the end of the Triassic period, the movement of depression recommenced in our area, though it was doubtless balanced by elevation elsewhere; modification and development, checked in the one province, went on in that "elsewhere";

and the chief forms of Mammals, Birds and Reptiles, as we know them, were evolved and peopled the Mesozoic continent.

I conceive Australia to have become separated from the continent as early as the end of the Triassic epoch, or not much later.

The Mesozoic continent must, I conceive, have lain to the east, about the shores of the North Pacific and Indian Oceans; and I am inclined to believe that it continued along the eastern side of the Pacific area to what is now the province of Austro-Columbia, the characteristic fauna of which is probably a remnant of the population of the latter part of this period.

Towards the latter part of the Mesozoic period the movement of upheaval around the shores of the Atlantic once more recommenced, and was very probably accompanied by a depression around those of the Pacific.

The Vertebrate fauna elaborated in the Mesozoic continent moved westward and took possession of the new lands, which gradually increased in extent up to, and in some directions after, the Miocene epoch.

It is in favour of this hypothesis, I think, that it is consistent with the persistence of a general uniformity in the positions of the great masses of land and water.

From the Devonian period, or earlier, to the present day, the four great oceans, Atlantic, Pacific, Arctic, and Antarctic, may have occupied their present positions, and only their coasts and channels of communication have undergone an incessant alteration.

And, finally, the hypothesis I have put before you requires no supposition that the rate of change in organic life has been either greater or less in ancient times than it is now; nor any assumption, either physical or biological, which has not its justification in analogous phenomena of existing nature.

I have now only to discharge the last duty of my office, which is to thank you, not only for the patient attention with which you have listened to me so long to-day, but also for the uniform kindness with which, for the past two years, you have rendered my endeavours to perform the important, and often laborious, functions of your President a pleasure instead of a burden.

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