The competition will generally be most severe, as formerly explained and illustrated by examples, between the forms which are most like each other in all respects. Hence the improved and modified descendants of a species will generally cause the extermination of the parent-species; and if many new forms have been developed from any one species, the nearest allies of that species, i.e. the species of the same genus, will be the most liable to extermination. Thus, as I believe, a number of new species descended from one species, that is a new genus, comes to supplant an old genus, belonging to the same family. But it must often have happened that a new species belonging to some one group will have seized on the place occupied by a species belonging to a distinct group, and thus caused its extermination; and if many allied forms be developed from the successful intruder, many will have to yield their places; and it will generally be allied forms, which will suffer from some inherited inferiority in common. But whether it be species belonging to the same or to a distinct class, which yield their places to other species which have been modified and improved, a few of the sufferers may often long be preserved, from being fitted to some peculiar line of life, or from inhabiting some distant and isolated station, where they have escaped severe competition. For instance, a single species of Trigonia, a great genus of shells in the secondary formations, survives in the Australian seas; and a few members of the great and almost extinct group of Ganoid fishes still inhabit our fresh waters. Therefore the utter extinction of a group is generally, as we have seen, a slower process than its production.
With respect to the apparently sudden extermination of whole families or orders, as of Trilobites at the close of the palæozoic period and of Ammonites at the close of the secondary period, we must remember what has been already said on the probable wide intervals of time between our consecutive formations; and in these intervals there may have been much slow extermination. Moreover, when by sudden immigration or by unusually rapid development, many species of a new group have taken possession of a new area, they will have exterminated in a correspondingly rapid manner many of the old inhabitants; and the forms which thus yield their places will commonly be allied, for they will partake of some inferiority in common.
Thus, as it seems to me, the manner in which single species and whole groups of species become extinct, accords well with the theory of natural selection. We need not marvel at extinction; if we must marvel, let it be at our presumption in imagining for a moment that we understand the many complex contingencies, on which the existence of each species depends. If we forget for an instant, that each species tends to increase inordinately, and that some check is always in action, yet seldom perceived by us, the whole economy of nature will be utterly obscured. Whenever we can precisely say why this species is more abundant in individuals than that; why this species and not another can be naturalised in a given country; then, and not till then, we may justly feel surprise why we cannot account for the extinction of this particular species or group of species.
On the Forms of Life changing almost simultaneously throughout the World.—Scarcely any palæontological discovery is more striking than the fact, that the forms of life change almost simultaneously throughout the world. Thus our European Chalk formation can be recognised in many distant parts of the world, under the most different climates, where not a fragment of the mineral chalk itself can be found; namely, in North America, in equatorial South America, in Tierra del Fuego, at the Cape of Good Hope, and in the peninsula of India. For at these distant points, the organic remains in certain beds present an unmistakeable degree of resemblance to those of the Chalk. It is not that the same species are met with; for in some cases not one species is identically the same, but they belong to the same families, genera, and sections of genera, and sometimes are similarly characterised in such trifling points as mere superficial sculpture. Moreover other forms, which are not found in the Chalk of Europe, but which occur in the formations either above or below, are similarly absent at these distant points of the world. In the several successive palæozoic formations of Russia, Western Europe and North America, a similar parallelism in the forms of life has been observed by several authors: so it is, according to Lyell, with the several European and North American tertiary deposits. Even if the few fossil species which are common to the Old and New Worlds be kept wholly out of view, the general parallelism in the successive forms of life, in the stages of the widely separated palæozoic and tertiary periods, would still be manifest, and the several formations could be easily correlated.
These observations, however, relate to the marine inhabitants of distant parts of the world: we have not sufficient data to judge whether the productions of the land and of fresh water change at distant points in the same parallel manner. We may doubt whether they have thus changed: if the Megatherium, Mylodon, Macrauchenia, and Toxodon had been brought to Europe from La Plata, without any information in regard to their geological position, no one would have suspected that they had coexisted with still living sea-shells; but as these anomalous monsters coexisted with the Mastodon and Horse, it might at least have been inferred that they had lived during one of the latter tertiary stages.
When the marine forms of life are spoken of as having changed simultaneously throughout the world, it must not be supposed that this expression relates to the same thousandth or hundred-thousandth year, or even that it has a very strict geological sense; for if all the marine animals which live at the present day in Europe, and all those that lived in Europe during the pleistocene period (an enormously remote period as measured by years, including the whole glacial epoch), were to be compared with those now living in South America or in Australia, the most skilful naturalist would hardly be able to say whether the existing or the pleistocene inhabitants of Europe resembled most closely those of the southern hemisphere. So, again, several highly competent observers believe that the existing productions of the United States are more closely related to those which lived in Europe during certain later tertiary stages, than to those which now live here; and if this be so, it is evident that fossiliferous beds deposited at the present day on the shores of North America would hereafter be liable to be classed with somewhat older European beds. Nevertheless, looking to a remotely future epoch, there can, I think, be little doubt that all the more modern marine formations, namely, the upper pliocene, the pleistocene and strictly modern beds, of Europe, North and South America, and Australia, from containing fossil remains in some degree allied, and from not including those forms which are only found in the older underlying deposits, would be correctly ranked as simultaneous in a geological sense.
The fact of the forms of life changing simultaneously, in the above large sense, at distant parts of the world, has greatly struck those admirable observers, MM. de Verneuil and d’Archiac. After referring to the parallelism of the palæozoic forms of life in various parts of Europe, they add, “If struck by this strange sequence, we turn our attention to North America, and there discover a series of analogous phenomena, it will appear certain that all these modifications of species, their extinction, and the introduction of new ones, cannot be owing to mere changes in marine currents or other causes more or less local and temporary, but depend on general laws which govern the whole animal kingdom.” M. Barrande has made forcible remarks to precisely the same effect. It is, indeed, quite futile to look to changes of currents, climate, or other physical conditions, as the cause of these great mutations in the forms of life throughout the world, under the most different climates. We must, as Barrande has remarked, look to some special law. We shall see this more clearly when we treat of the present distribution of organic beings, and find how slight is the relation between the physical conditions of various countries, and the nature of their inhabitants.
This great fact of the parallel succession of the forms of life throughout the world, is explicable on the theory of natural selection. New species are formed by new varieties arising, which have some advantage over older forms; and those forms, which are already dominant, or have some advantage over the other forms in their own country, would naturally oftenest give rise to new varieties or incipient species; for these latter must be victorious in a still higher degree in order to be preserved and to survive. We have distinct evidence on this head, in the plants which are dominant, that is, which are commonest in their own homes, and are most widely diffused, having produced the greatest number of new varieties. It is also natural that the dominant, varying, and far-spreading species, which already have invaded to a certain extent the territories of other species, should be those which would have the best chance of spreading still further, and of giving rise in new countries to new varieties and species. The process of diffusion may often be very slow, being dependent on climatal and geographical changes, or on strange accidents, but in the long run the dominant forms will generally succeed in spreading. The diffusion would, it is probable, be slower with the terrestrial inhabitants of distinct continents than with the marine inhabitants of the continuous sea. We might therefore expect to find, as we apparently do find, a less strict degree of parallel succession in the productions of the land than of the sea.
Dominant species spreading from any region might encounter still more dominant species, and then their triumphant course, or even their existence, would cease. We know not at all precisely what are all the conditions most favourable for the multiplication of new and dominant species; but we can, I think, clearly see that a number of individuals, from giving a better chance of the appearance of favourable variations, and that severe competition with many already existing forms, would be highly favourable, as would be the power of spreading into new territories. A certain amount of isolation, recurring at long intervals of time, would probably be also favourable, as before explained. One quarter of the world may have been most favourable for the production of new and dominant species on the land, and another for those in the waters of the sea. If two great regions had been for a long period favourably circumstanced in an equal degree, whenever their inhabitants met, the battle would be prolonged and severe; and some from one birthplace and some from the other might be victorious. But in the course of time, the forms dominant in the highest degree, wherever produced, would tend everywhere to prevail. As they prevailed, they would cause the extinction of other and inferior forms; and as these inferior forms would be allied in groups by inheritance, whole groups would tend slowly to disappear; though here and there a single member might long be enabled to survive.
Thus, as it seems to me, the parallel, and, taken in a large sense, simultaneous, succession of the same forms of life throughout the world, accords well with the principle of new species having been formed by dominant species spreading widely and varying; the new species thus produced being themselves dominant owing to inheritance, and to having already had some advantage over their parents or over other species; these again spreading, varying, and producing new species. The forms which are beaten and which yield their places to the new and victorious forms, will generally be allied in groups, from inheriting some inferiority in common; and therefore as new and improved groups spread throughout the world, old groups will disappear from the world; and the succession of forms in both ways will everywhere tend to correspond.