original genus, may for a long period continue to transmit unaltered descendants; and this is shown in the diagram by the dotted lines not prolonged far upwards from want of space.

But during the process of modification, represented in the diagram, another of our principles, namely that of extinction, will have played an important part. As in each fully stocked country natural selection necessarily acts by the selected form having some advantage in the struggle for life over other forms, there will be a constant tendency in the improved descendants of any one species to supplant and exterminate in each stage of descent their predecessors and their original parent. For it should be remembered that the competition will generally be most severe between those forms which are most nearly related to each other in habits, constitution, and structure. Hence all the intermediate forms between the earlier and later states, that is between the less and more improved states of a species, as well as the original parent-species itself, will generally tend to become extinct. So it probably will be with many whole collateral lines of descent, which will be conquered by later and improved lines of descent. If, however, the modified offspring of a species get into some distinct country, or become quickly adapted to some quite new station, in which child and parent do not come into competition, both may continue to exist.

If, then, our diagram be assumed to represent a considerable amount of modification, species (A) and all the earlier varieties will have become extinct, having been replaced by eight new species (a to m1); and (I) will have been replaced by six (n1 to z1) new species.

But we may go further than this. The original species of our genus were supposed to resemble each other in unequal degrees, as is so generally the case in nature; species (A) being more nearly related to B, C, and D, than to the other species; and species (I) more to G, H, K, L, than to the others. These two species (A) and (I), were also supposed to be very common and widely diffused species, so that they must originally have had some advantage over most of the other species of the genus. Their modified descendants, fourteen in number at the fourteen-thousandth generation, will probably have inherited some of the same advantages: they have also

been modified and improved in a diversified manner at each stage of descent, so as to have become adapted to many related places in the natural economy of their country. It seems, therefore, extremely probable that they will have taken the places of, and thus exterminated, not only their parents (A) and (I), but likewise some of the original species which were most nearly related to their parents. Hence very few of the original species will have transmitted offspring to the fourteen-thousandth generation. We may suppose that only one (F), of the two species which were least closely related to the other nine original species, has transmitted descendants to this late stage of descent.

The new species in our diagram, descended from the original eleven species, will now be fifteen in number. Owing to the divergent tendency of natural selection, the extreme amount of difference in character between species a1 and z1 will be much greater than that between the most different of the original eleven species. The new species, moreover, will be allied to each other in a widely different manner. Of the eight descendants from (A) the three marked a1, q1, p1, will be nearly related from having recently branched off from a10; b1 and f1, from having diverged at an earlier period from a, will be in some degree distinct from the three firstnamed species; and lastly, o1, e1⁄4, and m1, will be nearly related one to the other, but from having diverged at the first commencement of the process of modification, will be widely different from the other five species, and may constitute a sub-genus or even a distinct genus.

The six descendants from (I) will form two subgenera or even genera. But as the original species (I) differed largely from (A), standing nearly at the extreme points of the original genus, the six descendants from (I) will, owing to inheritance alone, differ considerably from the eight descendants from (A); the two groups, moreover, are supposed to have gone on diverging in different directions. The intermediate species, also (and this is a very important consideration), which connected the original species (A) and (I), have all become, excepting (F), extinct, and have left no descendants. Hence the six new species descended from (I), and the eight descended from (A), will have to be ranked as very distinct genera, or even as distinct sub-families.

Thus it is, as I believe, that two or more genera are produced by descent with modification, from two or more species of the same genus. And the two or more parentspecies are supposed to have descended from some one species of an earlier genus. In our diagram, this is indicated by the broken lines, beneath the capital letters, converging in sub-branches downwards towards a single point; this point representing a single species, the supposed single parent of our several new sub-genera and genera.

It is worth while to reflect for a moment on the character of the new species F1, which is supposed not to have diverged much in character, but to have retained the form of (F), either unaltered or altered only in a slight degree. In this case, its affinities to the other fourteen new species will be of a curious and circuitous nature. Having descended from a form which stood between the two parent species (A) and (I), now supposed to be extinct and unknown, it will be in some degree intermediate in character between the two groups descended from these species. But as these two groups have gone on diverging in character from the type of their parents, the new species (F) will not be directly intermediate between them, but rather between types of the two groups; and every naturalist will be able to bring some such case 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 hundred million generations, and likewise 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, or 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 very ancient 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 our diagram, we suppose the amount of change represented by each successive group of diverging dotted lines to be very great, the forms marked a1 to p1⁄4, those marked b1 and f1⁄44, and those marked o1 to m", will form

three very distinct genera. We shall also have two very distinct genera descended from (I); and as these latter two genera, both from continued divergence of character and from inheritance from a different parent, will differ widely from the three genera descended from (A), the two little groups of genera will form two distinct families, or even orders, according to the amount of divergent modification supposed to be represented in the diagram. And the two new families, or orders, will have descended from two species of the original genus; and these two species are supposed to have descended from one species of a still more ancient and unknown genus.

We have seen that in each country it is the species of 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 numbers, and thus lessen its chance of further variation and improvement. Within the same large group, the later and more highly perfected sub-groups, 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 sub-groups. Small and broken groups and sub-groups 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 as yet have suffered least extinction, will for a long period continue to increase. But which groups

will ultimately prevail, no man can predict; for we well 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 on this view of extremely few of the more ancient species having transmitted descendants, and on the view of all the descendants of the same species making a class, we can understand how it is that there exist but very few classes in each main division of the animal and vegetable kingdoms. Although extremely few of the most ancient species now have living and modified descendants, yet at the most remote geological period, the earth may have been as well peopled with many 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, as we have seen, exclusively by the preservation and accumulation of variations, which are beneficial under the organic and inorganic conditions of life to which each creature is at each successive period exposed. The ultimate result will be that each creature will tend to become more and more improved in relation to its conditions of life. This improvement will, I think, inevitably lead 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. Amongst 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 undergo in their development from the 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 different parts (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 a subject this is if we look, for instance, to fish, amongst which some naturalists rank those as highest which, like the sharks, approach nearest to reptiles; whilst other naturalists rank the