Origin of Species Variorum

Comparison with 1869

methodical selection, or by .. unconscious selection which results from each man trying to keep the best dogs without any thought of modifying the breed. I may add, that, according to Mr. Pierce, there are two varieties of the wolf inhabiting the Catskill Mountains in the United States, one with a light greyhound-like form, which pursues deer, and the other more bulky, with shorter legs, which more frequently attacks the shepherd's flocks.

It should be observed that, in the above illustration, I speak of the slimmest individual wolves, and not of any single strongly-marked variation having been preserved. In former editions of this work I sometimes spoke as if this latter alternative had frequently occurred. I saw the great importance of individual differences, and this led me fully to discuss the results of unconscious selection by man, which depends on the preservation of the better adapted or more valuable individuals, and on the destruction of the worst. I saw, also, that the preservation in a state of nature of any occasional deviation of structure, such as a monstrosity, would be a rare event; and that, if preserved, it would generally be lost by subsequent intercrossing with ordinary individuals. Nevertheless, until reading an able and valuable article in the 'North British Review' (1867), I did not appreciate how rarely single variations, whether slight or strongly-marked, could be perpetuated. The author takes the case of a pair of animals, which produce during their lifetime two hundred offspring, of which, from various causes of destruction, only two on an average survive to procreate their kind. This is rather an extreme estimate for most of the higher animals, but by no means so for many of the lower organisms. He then shows that if a single individual were born, which varied in some manner, giving it twice as good a chance of life as that of the other individuals, yet the chances would be strongly against its survival. Supposing it to survive and to breed, and that half its young inherited the favourable variation; still, as the Reviewer goes on to show, the young would have only a slightly better chance of surviving and breeding; and this chance would go on decreasing in the succeeding generations. The justice of these remarks cannot, I think, be disputed. If, for instance, a bird of some kind could procure its food more easily by having its beak curved, and if one were born with its beak strongly curved, and which consequently flourished, nevertheless there would be a very poor chance of this one individual perpetuating its kind to the exclusion of the common form; but there can hardly be a doubt, judging by what we see taking place under domestication, that this result would follow from the preservation during many generations of a large number of individuals with more or less curved beaks, and from the destruction of a still larger number with the straightest beaks.

It should not, however, be overlooked that certain variations, which no one would rank as mere individual differences, frequently recur owing to a similar organisation being similarly acted on,—of which fact numerous instances could be given with our domestic productions. In such cases, if a varying individual did not actually transmit to its offspring its newly-acquired character, it would undoubtedly transmit, as long as the existing conditions remained the same, a still stronger tendency to vary in the same manner. The conditions might indeed act in so energetic and definite a manner as to lead to the same modification in all the individuals of the species without the aid of selection. ↑ But we may suppose that the conditions sufficed to affect only a third, or fourth, or tenth part of the individuals; and several such cases could be given; for instance, it has been estimated by Graba that in the Faroe Islands about one-fifth of the guillemots, which all breed together, consist of a well-marked variety; and this was formerly ranked as a distinct species under the name of Uria lacrymans. Now, in such cases, if the variation were of a beneficial nature, the original form would soon be supplanted by the modified form, through the survival of the fittest.

With reference to the effects of intercrossing and of competition, it should be borne in mind that most animals and plants keep to their proper homes, and do not needlessly wander about; we see this even with migratory birds, which almost always return to the same district. Consequently each newly-formed variety would generally be at first local, as seems to be the common rule with varieties in a state of nature; so that similarly modified individuals would soon exist in a small body together, and would often breed together. If the new variety was successful in its battle for life, it would slowly spread from a central spot, competing with and conquering the unchanged individuals on the margins of an ever-increasing circle. But to the subject of intercrossing we shall have to return. It may be objected by those who have not attended to natural history, that the long-continued accumulation of individual differences could not give rise to parts or organs which seem to us, and are often called, new. But, as we shall hereafter find, it is difficult to advance any good instance of a really new organ; even so complex and perfect an organ as the eye can be shown to graduate downwards into mere tissue sensitive to diffused light.

It may be worth while to give another and more complex illustration of the action of natural selection. ↑ Certain plants excrete .. sweet juice, apparently for the sake of eliminating

methodical selection, or by that unconscious selection which results from each man trying to keep the best dogs without any thought of modifying the breed. ↑

14 blocks not present in 1859 1860 1861 1866; present in 1869 1872

I may add, that, according to Mr. Pierce, there are two varieties of the wolf inhabiting the Catskill Mountains in the United States, one with a light greyhound-like form, which pursues deer, and the other more bulky, with shorter legs, which more frequently attacks the shepherd's flocks. It should be observed that, in the above illustration, I speak of the slimmest individual wolves, and not of any single strongly-marked variation having been preserved. In former editions of this work I sometimes spoke as if this latter alternative had frequently occurred. I saw the great importance of individual differences, and this led me fully to discuss the results of unconscious selection by man, which depends on the preservation of the better adapted or more valuable individuals, and on the destruction of the worst. I saw, also, that the preservation in a state of nature of any occasional deviation of structure, such as a monstrosity, would be a rare event; and that, if preserved, it would generally be lost by subsequent intercrossing with ordinary individuals. Nevertheless, until reading an able and valuable article in the 'North British Review' (1867), I did not appreciate how rarely single variations, whether slight or strongly-marked, could be perpetuated. The author takes the case of a pair of animals, which produce during their lifetime two hundred offspring, of which, from various causes of destruction, only two on an average survive to procreate their kind. This is rather an extreme estimate for most of the higher animals, but by no means so for many of the lower organisms. He then shows that if a single individual were born, which varied in some manner, giving it twice as good a chance of life as that of the other individuals, yet the chances would be strongly against its survival. Supposing it to survive and to breed, and that half its young inherited the favourable variation; still, as the Reviewer goes on to show, the young would have only a slightly better chance of surviving and breeding; and this chance would go on decreasing in the succeeding generations. The justice of these remarks cannot, I think, be disputed. If, for instance, a bird of some kind could procure its food more easily by having its beak curved, and if one were born with its beak strongly curved, and which consequently flourished, nevertheless there would be a very poor chance of this one individual perpetuating its kind to the exclusion of the common form; but there can hardly be a doubt, judging by what we see taking place under domestication, that this result would follow from the preservation during many generations of a large number of individuals with more or less curved beaks, and from the destruction of a still larger number with the straightest beaks. It should not, however, be overlooked that certain variations, which no one would rank as mere individual differences, frequently recur owing to a similar organisation being similarly acted on,—of which fact numerous instances could be given with our domestic productions. In such cases, if a varying individual did not actually transmit to its offspring its newly-acquired character, it would undoubtedly transmit, as long as the existing conditions remained the same, a still stronger tendency to vary in the same manner.

↑

Even without any change in the proportional numbers of the animals on which our wolf preyed, a cub might be born with an innate tendency to pursue certain kinds of prey. Nor can this be thought very improbable; for we often observe great differences in the natural tendencies of our domestic animals; one cat, for instance, taking to catch rats, another mice; one cat, according to Mr. St. John, bringing home winged game, another hares or rabbits, and another hunting on marshy ground and almost nightly catching woodcocks or snipes. The tendency to catch rats rather than mice is known to be inherited. Now, if any slight innate change of habit or of structure benefited an individual wolf, it would have the best chance of surviving and of leaving offspring. Some of its young would probably inherit the same habits or structure, and by the repetition of this process, a new variety might be formed which would either supplant, or coexist with, the parent form of .. wolf. Or, again, the wolves inhabiting a mountainous district, and those frequenting the lowlands, would naturally be forced to hunt different prey; and from the continued preservation of the individuals best fitted for the two sites, two varieties would slowly be formed. These varieties would cross and blend where they met; but to this subject of intercrossing we shall soon have to return. I may add, that, according to Mr. Pierce, there are two varieties of the wolf inhabiting the Catskill Mountains in the United States, one with a light greyhound-like form, which pursues deer, and the other more bulky, with shorter legs, which more frequently attacks the shepher's flocks.

Let us now take a more complex case. Certain plants excrete a sweet juice, apparently for the sake of elimi- nating