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or flesh alone, draws most nutriment from these substances. So in the general economy of any land, the more widely and perfectly the animals and plants are diversified for different habits of life, so will a greater number of individuals be capable of there supporting themselves. A set of animals, with their organisation but little diversified, could hardly compete with a set more perfectly diversified in structure. It may be doubted, for instance, whether the Australian marsupials, which are divided into groups differing but little from each other, and feebly representing, as Mr. Waterhouse and others have remarked, our carnivorous, ruminant, and rodent mammals, could successfully compete with these well-developed orders. In the Australian mammals, we see the process of diversification in an early and incomplete stage of development.
The Probable Effects of the Action of Natural Selection through Divergence of Character and Extinction, on the Descendants of a Common Ancestor.
After the foregoing discussion, which has been much compressed, we may assume that the modified descendants of any one species will succeed so much the better as they become more diversified in structure, and are thus enabled to encroach on places occupied by other beings. Now let us see how this principle of benefit being derived from divergence of character, combined with the principles of natural selection and of extinction, tends to act.
The accompanying diagram will aid us in understanding this rather perplexing subject. Let A to L represent the species of a genus large in its own country; these species are supposed to resemble each other in unequal degrees, as is so generally the case in nature, and as is represented in the diagram by the letters standing at unequal distances. I have said a large genus, because as we saw in the second chapter, on an average more species vary in large genera than in small genera; and the varying species of the large genera present a greater number of varieties. We have, also, seen that the species, which are the commonest and the most widely diffused, vary more than do the rare and restricted species. Let (A) be a common, widely-diffused, and varying species, belonging to a genus large in its own country. The branching and diverging dotted lines of unequal lengths proceeding from (A), may represent its varying offspring. The variations are supposed to be extremely slight, but of the most diversified nature; they are not supposed all to appear simultaneously, but often after long intervals of time; nor are they all supposed to endure for equal periods. Only those variations which are in some way profitable will be preserved or naturally selected. And here the importance of the principle of benefit derived from divergence of character comes in; for this will generally lead to the most different or divergent variations (represented by the outer dotted lines) being preserved and accumulated by natural selection. When a dotted line reaches one of the horizontal lines, and is there marked by a small numbered letter, a sufficient amount of variation is supposed to have been accumulated to form it into a fairly well-marked variety, such as would be thought worthy of record in a systematic work. The intervals between the horizontal lines in the diagram, may represent each a thousand or more generations. After a thousand generations, species (A) is supposed to have produced two fairly well-marked varieties, namely a' and mo. These two varieties will generally still be exposed to the same conditions which made their parents variable, and the tendency to variability is in itself hereditary; consequently they will likewise tend to vary, and commonly in nearly the same manner as did their parents. Moreover, these two varieties, being only slightly modified forms, will tend to inherit those advantages which made their parent (A) more numerous than most of the other inhabitants of the same country; they will also partake of those more general advantages which made the genus to which the parent-species belonged, a large genus in its own country. And all these circumstances are favourable to the production of new varieties. If, then, these two varieties be variable, the most divergent of their variations will generally be preserved during the next thousand generations. And after this interval, variety a' is supposed in the diagram to have produced variety a”, which will, owing to the principle of divergence, differ more from (A) than did variety a". Variety m' is supposed to have produced two varieties, namely m” and so, differing from each other, and more considerably from their common parent (A). We may continue the process by similar steps for any length of time; some of the varieties, after each thousand generations, producing only a single variety, but in a more and more modified condition, some producing two or three varieties, and some failing to produce any. Thus the varieties or modified descendants of the common parent (A), will generally go on increasing in number and diverging in character. In the diagram the process is represented up to the ten-thousandth generation, and under a condensed and simplified form up to the fourteen-thousandth generation. But I must here remark that I do not suppose that the process ever goes on so regularly as is represented in the diagram, though in itself made somewhat irregular, nor that it goes on continuously; it is far more probable that each form remains for long periods unaltered, and then again undergoes modification, Nor do I
suppose that the most divergent varieties are invariably preserved: a medium form may often long endure, and may or may not produce more than one modified descendant; for natural selection will always act according to the nature of the places which are either unoccupied or not perfectly occupied by other beings; and this will depend on infinitely complex relations. But as a general rule, the more diversified in structure the descendants from any one species can be rendered, the more places they will be enabled to seize on, and the more their modified progeny will increase. In our diagram the line of succession is broken at regular intervals by small numbered letters marking the successive forms which have become sufficiently distinct to be recorded as varieties. But these breaks are imaginary, and might have been inserted anywhere, after intervals long enough to allow the accumulation of a considerable amount of divergent variation. As all the modified descendants from a common and widelydiffused species, belonging to a large genus, will tend to partake of the same advantages which made their parent successful in life, they will generally go on multiplying in number as well as diverging in character: this is represented in the diagram by the several divergent branches proceeding from (A). The modified offspring from the later and more highly improved branches in the lines of descent, will, it is probable, often take the place of, and so destroy, the earlier and less improved branches: this is represented in the diagram by some of the lower branches not reaching to the upper horizontal lines. In some cases no doubt the process of modification will be confined to a single line of descent, and the number of modified descendants will not be increased; although the amount of divergent modification may have been augmented. This case would be represented in the diagram, if all the lines proceeding from (A) were removed, excepting that from a to al". In the same way the English race-horse and English pointer have apparently both gone on slowly diverging in character from their original stocks, without either having given off any fresh branches Or races. After ten thousand generations, species (A) is supposed to have produced three forms, a”, f", and m”, which, from having diverged in character during the successive generations, will have come to differ largely, but perhaps unequally, from each other and from their common parent. If we suppose the amount of change between each horizontal line in our diagram to be excessively small, these three forms may still be only well-marked varieties; but we have only to suppose the steps in the process of modification to be more numerous or greater in amount, to convert these three forms into doubtful or at least into well-defined species. Thus the diagram illustrates the steps by which the small differences distinguishing varieties are increased into the larger differences distinguishing species. By continuing the same process for a greater number of generations (as shown in the diagram in a condensed and simplified manner), we get eight species, marked by the letters between a" and mo, all descended from (A). Thus, as I believe, species are multiplied and genera are formed. In a large genus it is probable that more than one species would vary. In the diagram I have assumed that a second species (I) has produced, by analogous steps, after ten thousand generations, either two well-marked varieties (w" and z") or two species, according to the amount of change supposed to be represented between the horizontal lines. After fourteen thousand generations, six new species, marked by the letters no to z*, are supposed to have been produced. In any genus, the species which are already very different in character from each other, will generally tend to produce the greatest number of modified descendants; for these will have the best chance of seizing on new and widely different places in the polity of nature: hence in the diagram I have chosen the extreme species (A), and the nearly extreme species (I), as those which have largely varied, and have given rise to new varieties and species. The other nine species (marked by capital letters) of our original genus, may for long but unequal periods continue to transmit unaltered descendants; and this is shown in the diagram by the dotted lines unequally prolonged upwards. 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 progenitor. 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 the same 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. If, however, the modified offspring of a species get into some distinct country, or become quickly adapted to some quite new station, in which offspring and progenitor 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, being replaced by eight new species (a” to mo); and species (I) will be replaced by six (no to z*) 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 fourteenthousandth 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 (E and F) 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 a” and z* will be much greater than that between the most distinct 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 a”, g”, p", will be nearly related from having recently branched off from a "; bo. and f*, from having diverged at an earlier period from a”, will be in some degree distinct from the three first-named species; and lastly, oo, e”, and mo, 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 subgenus or a distinct genus. The six descendants from (I) will form two sub-genera or genera. But as the original species (I) differed largely from (A), standing nearly at the extreme end 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