But let the external conditions of a country change; if in a small degree, the relative proportions of the inhabitants will in most cases simply be slightly changed; but let the number of inhabitants be small, as in an island[{232}], and free access to it from other countries be circumscribed; and let the change of condition continue progressing (forming new stations); in such case the original inhabitants must cease to be so perfectly adapted to the changed conditions as they originally were. It has been shown that probably such changes of external conditions would, from acting on the reproductive system, cause the organization of the beings most affected to become, as under domestication, plastic. Now can it be doubted from the struggle each individual (or its parents) has to obtain subsistence that any minute variation in structure, habits, or instincts, adapting that individual better to the new conditions, would tell upon its vigour and health? In the struggle it would have a better chance of surviving, and those of its offspring which inherited the variation, let it be ever so slight, would have a better chance to survive. Yearly more are bred than can survive; the smallest grain in the balance, in the long run, must tell on which death shall fall, and which shall survive[{233}]. Let this work of selection, on the one hand, and death on the other, go on for a thousand generations; who would pretend to affirm that it would produce no effect, when we remember what in a few years Bakewell effected in cattle and Western in sheep, by this identical principle of selection.
To give an imaginary example, from changes in progress on an island, let the organization[{234}] of a canine animal become slightly plastic, which animal preyed chiefly on rabbits, but sometimes on hares; let these same changes cause the number of rabbits very slowly to decrease and the number of hares to increase; the effect of this would be that the fox or dog would be driven to try to catch more hares, and his numbers would tend to decrease; his organization, however, being slightly plastic, those individuals with the lightest forms, longest limbs, and best eye-sight (though perhaps with less cunning or scent) would be slightly favoured, let the difference be ever so small, and would tend to live longer and to survive during that time of the year when food was shortest; they would also rear more young, which young would tend to inherit these slight peculiarities. The less fleet ones would be rigidly destroyed. I can see no more reason to doubt but that these causes in a thousand generations would produce a marked effect, and adapt the form of the fox to catching hares instead of rabbits, than that greyhounds can be improved by selection and careful breeding. So would it be with plants under similar circumstances; if the number of individuals of a species with plumed seeds could be increased by greater powers of dissemination within its own area (that is if the check to increase fell chiefly on the seeds), those seeds which were provided with ever so little more down, or with a plume placed so as to be slightly more acted on by the winds, would in the long run tend to be most disseminated; and hence a greater number of seeds thus formed would germinate, and would tend to produce plants inheriting this slightly better adapted down.
Besides this natural means of selection, by which those individuals are preserved, whether in their egg or seed or in their mature state, which are best adapted to the place they fill in nature, there is a second agency at work in most bisexual animals tending to produce the same effect, namely the struggle of the males for the females. These struggles are generally decided by the law of battle; but in the case of birds, apparently, by the charms of their song[{235}], by their beauty or their power of courtship, as in the dancing rock-thrush of Guiana. Even in the animals which pair there seems to be an excess of males which would aid in causing a struggle: in the polygamous animals[{236}], however, as in deer, oxen, poultry, we might expect there would be severest struggle: is it not in the polygamous animals that the males are best formed for mutual war? The most vigorous males, implying perfect adaptation, must generally gain the victory in their several contests. This kind of selection, however, is less rigorous than the other; it does not require the death of the less successful, but gives to them fewer descendants. This struggle falls, moreover, at a time of year when food is generally abundant, and perhaps the effect chiefly produced would be the alteration of sexual characters, and the selection of individual forms, no way related to their power of obtaining food, or of defending themselves from their natural enemies, but of fighting one with another. This natural struggle amongst the males may be compared in effect, but in a less degree, to that produced by those agriculturalists who pay less attention to the careful selection of all the young animals which they breed and more to the occasional use of a choice male[{237}].
Differences between “Races” and “Species”:—first, in their trueness or variability.
Races[{238}] produced by these natural means of selection[{239}] we may expect would differ in some respects from those produced by man. Man selects chiefly by the eye, and is not able to perceive the course of every vessel and nerve, or the form of the bones, or whether the internal structure corresponds to the outside shape. He[{240}] is unable to select shades of constitutional differences, and by the protection he affords and his endeavours to keep his property alive, in whatever country he lives, he checks, as much as lies in his power, the selecting action of nature, which will, however, go on to a lesser degree with all living things, even if their length of life is not determined by their own powers of endurance. He has bad judgment, is capricious, he does not, or his successors do not, wish to select for the same exact end for hundreds of generations. He cannot always suit the selected form to the properest conditions; nor does he keep those conditions uniform: he selects that which is useful to him, not that best adapted to those conditions in which each variety is placed by him: he selects a small dog, but feeds it highly; he selects a long-backed dog, but does not exercise it in any peculiar manner, at least not during every generation. He seldom allows the most vigorous males to struggle for themselves and propagate, but picks out such as he possesses, or such as he prefers, and not necessarily those best adapted to the existing conditions. Every agriculturalist and breeder knows how difficult it is to prevent an occasional cross with another breed. He often grudges to destroy an individual which departs considerably from the required type. He often begins his selection by a form or sport considerably departing from the parent form. Very differently does the natural law of selection act; the varieties selected differ only slightly from the parent forms[{241}]; the conditions are constant for long periods and change slowly; rarely can there be a cross; the selection is rigid and unfailing, and continued through many generations; a selection can never be made without the form be better adapted to the conditions than the parent form; the selecting power goes on without caprice, and steadily for thousands of years adapting the form to these conditions. The selecting power is not deceived by external appearances, it tries the being during its whole life; and if less well «?» adapted than its congeners, without fail it is destroyed; every part of its structure is thus scrutinised and proved good towards the place in nature which it occupies.
We have every reason to believe that in proportion to the number of generations that a domestic race is kept free from crosses, and to the care employed in continued steady selection with one end in view, and to the care in not placing the variety in conditions unsuited to it; in such proportion does the new race become “true” or subject to little variation[{242}]. How incomparably “truer” then would a race produced by the above rigid, steady, natural means of selection, excellently trained and perfectly adapted to its conditions, free from stains of blood or crosses, and continued during thousands of years, be compared with one produced by the feeble, capricious, misdirected and ill-adapted selection of man. Those races of domestic animals produced by savages, partly by the inevitable conditions of their life, and partly unintentionally by their greater care of the individuals most valuable to them, would probably approach closest to the character of a species; and I believe this is the case. Now the characteristic mark of a species, next, if not equal in importance to its sterility when crossed with another species, and indeed almost the only other character (without we beg the question and affirm the essence of a species, is its not having descended from a parent common to any other form), is the similarity of the individuals composing the species, or in the language of agriculturalists their “trueness.”
Difference between “Races” and “Species” in fertility when crossed.
The sterility of species, or of their offspring, when crossed has, however, received more attention than the uniformity in character of the individuals composing the species. It is exceedingly natural that such sterility[{243}] should have been long thought the certain characteristic of species. For it is obvious that if the allied different forms which we meet with in the same country could cross together, instead of finding a number of distinct species, we should have a confused and blending series. The fact however of a perfect gradation in the degree of sterility between species, and the circumstance of some species most closely allied (for instance many species of crocus and European heaths) refusing to breed together, whereas other species, widely different, and even belonging to distinct genera, as the fowl and the peacock, pheasant and grouse[{244}], Azalea and Rhododendron, Thuja and Juniperus, breeding together ought to have caused a doubt whether the sterility did not depend on other causes, distinct from a law, coincident with their creation. I may here remark that the fact whether one species will or will not breed with another is far less important than the sterility of the offspring when produced; for even some domestic races differ so greatly in size (as the great stag-greyhound and lap-dog, or cart-horse and Burmese ponies) that union is nearly impossible; and what is less generally known is, that in plants Kölreuter has shown by hundreds of experiments that the pollen of one species will fecundate the germen of another species, whereas the pollen of this latter will never act on the germen of the former; so that the simple fact of mutual impregnation certainly has no relation whatever to the distinctness in creation of the two forms. When two species are attempted to be crossed which are so distantly allied that offspring are never produced, it has been observed in some cases that the pollen commences its proper action by exserting its tube, and the germen commences swelling, though soon afterwards it decays. In the next stage in the series, hybrid offspring are produced though only rarely and few in number, and these are absolutely sterile: then we have hybrid offspring more numerous, and occasionally, though very rarely, breeding with either parent, as is the case with the common mule. Again, other hybrids, though infertile inter se, will breed quite freely with either parent, or with a third species, and will yield offspring generally infertile, but sometimes fertile; and these latter again will breed with either parent, or with a third or fourth species: thus Kölreuter blended together many forms. Lastly it is now admitted by those botanists who have longest contended against the admission, that in certain families the hybrid offspring of many of the species are sometimes perfectly fertile in the first generation when bred together: indeed in some few cases Mr Herbert[{245}] found that the hybrids were decidedly more fertile than either of their pure parents. There is no way to escape from the admission that the hybrids from some species of plants are fertile, except by declaring that no form shall be considered as a species, if it produces with another species fertile offspring: but this is begging the question[{246}]. It has often been stated that different species of animals have a sexual repugnance towards each other; I can find no evidence of this; it appears as if they merely did not excite each others passions. I do not believe that in this respect there is any essential distinction between animals and plants; and in the latter there cannot be a feeling of repugnance.