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to be extinct and unknown, it will be in some degree intermediate in character between the two groups descended from these two species. But as these two groups have gone on diverging in character from the type of their parents, the new species (F14) will not be directly intermediate between them, but rather between types of the two groups; and every naturalist will be able to call such cases before his mind.

In the diagram each horizontal line has hitherto been supposed to represent a thousand generations, but each may represent a million or more generations; it may also represent a section of the successive strata of the earth’s crust including extinct remains. We shall, when we come to our chapter on geology, have to refer again to this subject, and I think we shall then see that the diagram throws light on the affinities of extinct beings, which, though generally belonging to the same orders, families, or genera, with those now living, yet are often, in some degree, intermediate in character between existing groups; and we can understand this fact, for the extinct species lived at various remote epochs when the branching lines of descent had diverged less.

I see no reason to limit the process of modification, as now explained, to the formation of genera alone. If, in the diagram, we suppose the amount of change represented by each successive group of diverging dotted lines to be great, the forms marked a14 to p14, those marked b14 and f14, and those marked o14 to m14, will form three very distinct genera. We shall also have two very distinct genera descended from (I), differing widely from the descendants of (A). These two groups of genera will thus form two distinct families, or orders, according to the amount of divergent modification supposed to be represented in the diagram. And the two new families, or orders, are descended from two species of the original genus; and these are supposed to be descended from some still more ancient and unknown form.

We have seen that in each country it is the species belonging to the larger genera which oftenest present varieties or incipient species. This, indeed, might have been expected; for as natural selection acts through one form having some advantage over other forms in the struggle for existence, it will chiefly act on those which already have some advantage; and the largeness of any group shows that its species have inherited from a common ancestor some advantage in common. Hence, the struggle for the production of new and modified descendants will mainly lie between the larger groups, which are all trying to increase in number. One large group will slowly conquer another large group, reduce its number, and thus lessen its chance of further variation and improvement. Within the same large group, the later and more highly perfected subgroups, from branching out and seizing on many new places in the polity of nature, will constantly tend to supplant and destroy the earlier and less improved subgroups. Small and broken groups and subgroups will finally disappear. Looking to the future, we can predict that the groups of organic beings which are now large and triumphant, and which are least broken up, that is, which have as yet suffered least extinction, will, for a long period, continue to increase. But which groups will ultimately prevail, no man can predict; for we know that many groups, formerly most extensively developed, have now become extinct. Looking still more remotely to the future, we may predict that, owing to the continued and steady increase of the larger groups, a multitude of smaller groups will become utterly extinct, and leave no modified descendants; and consequently that, of the species living at any one period, extremely few will transmit descendants to a remote futurity.

I shall have to return to this subject in the chapter on classification, but I may add that as, according to this view, extremely few of the more ancient species have transmitted descendants to the present day, and, as all the descendants of the same species form a class, we can understand how it is that there exist so few classes in each main division of the animal and vegetable kingdoms. Although few of the most ancient species have left modified descendants, yet, at remote geological periods, the earth may have been almost as well peopled with species of many genera, families, orders and classes, as at the present day.

ON THE DEGREE TO WHICH ORGANISATION TENDS TO ADVANCE.

Natural selection acts exclusively by the preservation and accumulation of variations, which are beneficial under the organic and inorganic conditions to which each creature is exposed at all periods of life. The ultimate result is that each creature tends to become more and more improved in relation to its conditions. This improvement inevitably leads to the gradual advancement of the organisation of the greater number of living beings throughout the world. But here we enter on a very intricate subject, for naturalists have not defined to each other’s satisfaction what is meant by an advance in organisation. Among the vertebrata the degree of intellect and an approach in structure to man clearly come into play. It might be thought that the amount of change which the various parts and organs pass through in their development from embryo to maturity would suffice as a standard of comparison; but there are cases, as with certain parasitic crustaceans, in which several parts of the structure become less perfect, so that the mature animal cannot be called higher than its larva.

Von Baer’s standard seems the most widely applicable and the best, namely, the amount of differentiation of the parts of the same organic being, in the adult state, as I should be inclined to add, and their specialisation for different functions; or, as Milne Edwards would express it, the completeness of the division of physiological labour. But we shall see how obscure this subject is if we look, for instance, to fishes, among which some naturalists rank those as highest which, like the sharks, approach nearest to amphibians; while other naturalists rank the common bony or teleostean fishes as the highest, inasmuch as they are most strictly fish-like, and differ most from the other vertebrate classes. We see still more plainly the obscurity of the subject by turning to plants, among which the standard of intellect is of course quite excluded; and here some botanists rank those plants as highest which have every organ, as sepals, petals, stamens and pistils, fully developed in each flower; whereas other botanists, probably with more truth, look at the plants which have their several organs much modified and reduced in number as the highest.

If we take as the standard of high organisation, the amount of differentiation and specialisation of the several organs in each being when adult (and this will include the advancement of the brain for intellectual purposes), natural selection clearly leads towards this standard: for all physiologists admit that the specialisation of organs, inasmuch as in this state they perform their functions better, is an advantage to each being; and hence the accumulation of variations tending towards specialisation is within the scope of natural selection. On the other hand, we can see, bearing in mind that all organic beings are striving to increase at a high ratio and to seize on every unoccupied or less well occupied place in the economy of nature, that it is quite possible for natural selection gradually to fit a being to a situation in which several organs would be superfluous or useless: in such cases there would be retrogression in the scale of organisation. Whether organisation on the whole has actually advanced from the remotest geological periods to the present day will be more conveniently discussed in our chapter on Geological Succession.

But it may be objected that if all organic beings thus tend to rise in the scale, how is it that throughout the world a multitude of the lowest forms still exist; and how is it that in each great class some forms are far more highly developed than others? Why have not the more highly developed forms every where supplanted and exterminated the lower? Lamarck, who believed in an innate and inevitable tendency towards perfection in all organic beings, seems to have felt this difficulty so strongly that he was led to suppose that new and simple forms are continually being produced by spontaneous generation. Science has not as yet proved the truth of this belief, whatever the future may reveal. On our theory the continued existence of lowly organisms offers no difficulty; for natural selection, or the survival of the fittest, does not necessarily include progressive development—it only takes advantage of such variations as arise and are beneficial to each creature under its complex relations of life. And it may be asked what advantage, as far as we can see, would it be to an infusorian animalcule—to an intestinal worm—or even to an earth-worm, to be highly organised. If it were no advantage, these forms would be left, by natural selection, unimproved or but little improved, and might remain for indefinite ages in their present lowly condition. And geology tells us that some of the lowest forms, as the infusoria and rhizopods, have remained for an enormous period in nearly their present state. But to suppose that most of the many now existing low forms have not in the least advanced since the first dawn of life would be extremely rash; for every naturalist who has dissected some of the beings now ranked as very low in the scale, must have been struck with their really wondrous and beautiful organisation.

Nearly the same remarks are applicable, if we look to the different grades of organisation within the same great group; for instance, in the vertebrata, to the coexistence of mammals and fish—among mammalia, to the coexistence of man and the ornithorhynchus—among fishes, to the coexistence of the shark and the lancelet (Amphioxus), which latter fish in the extreme simplicity of its structure approaches the invertebrate classes. But mammals and fish hardly come into competition with each other; the advancement of the whole class of mammals, or of certain members in this class, to the highest grade would not lead to their taking the place of fishes. Physiologists believe that the brain must be bathed by warm blood to be highly active, and this requires aerial respiration; so that warm-blooded mammals when inhabiting the water lie under a disadvantage in having to come continually to the surface to breathe. With fishes, members of the shark family would not tend to supplant the lancelet; for the lancelet, as I hear from Fritz Muller, has as sole companion and competitor on the barren sandy shore of South Brazil, an anomalous annelid. The three lowest orders of mammals, namely, marsupials, edentata, and rodents, coexist in South America in the same region with numerous monkeys, and probably interfere little with each other. Although organisation, on the whole, may have advanced and be still advancing throughout the world, yet the scale will always present many degrees of perfection; for the high advancement of certain whole classes, or of certain members of each class, does not at all necessarily lead to the extinction of those groups with which they do not enter into close competition. In some cases, as we shall hereafter see, lowly organised forms appear to have been preserved to the present day, from inhabiting confined or peculiar stations, where they have been subjected to less severe competition, and where their scanty numbers have retarded the chance of favourable variations arising.

Finally, I believe that many lowly organised forms now exist throughout the world, from various causes. In some cases variations or individual differences of a favourable nature may never have arisen for natural selection to act on and accumulate. In no case, probably, has time sufficed for the utmost possible amount of development.

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