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.