Darwin and Modern Science [24]
grass and plants, have been evolved? And finally, how is it that the same Hawk-moth caterpillars, which to-day show oblique stripes, possessed longitudinal stripes in Tertiary times? We can read this fact from the history of their development, and I have before attempted to show the biological significance of this change of colour. ("Studien zur Descendenz-Theorie" II., "Die Enstehung der Zeichnung bei den Schmetterlings-raupen," Leipzig, 1876.)
For the present I need only draw the conclusion that one and the same caterpillar may exhibit the initial stages of both, and that it depends on the manner in which these marking elements are INTENSIFIED and COMBINED by natural selection whether whitish longitudinal or oblique stripes should result. In this case then the "useful variations" were actually "always there," and we see that in the same group of Lepidoptera, e.g. species of Sphingidae, evolution has occurred in both directions according to whether the form lived among grass or on broad leaves with oblique lateral veins, and we can observe even now that the species with oblique stripes have longitudinal stripes when young, that is to say, while the stripes have no biological significance. The white places in the skin which gave rise, probably first as small spots, to this protective marking could be combined in one way or another according to the requirements of the species. They must therefore either have possessed selection-value from the first, or, if this was not the case at their earliest occurrence, there must have been SOME OTHER FACTORS which raised them to the point of selection-value. I shall return to this in discussing germinal selection. But the case may be followed still farther, and leads us to the same alternative on a still more secure basis.
Many years ago I observed in caterpillars of Smerinthus populi (the poplar hawk-moth), which also possess white oblique stripes, that certain individuals showed RED SPOTS above these stripes; these spots occurred only on certain segments, and never flowed together to form continuous stripes. In another species (Smerinthus tiliae) similar blood-red spots unite to form a line-like coloured seam in the last stage of larval life, while in S. ocellata rust-red spots appear in individual caterpillars, but more rarely than in S. Populi, and they show no tendency to flow together.
Thus we have here the origin of a new character, arising from small beginnings, at least in S. tiliae, in which species the coloured stripes are a normal specific character. In the other species, S. populi and S. ocellata, we find the beginnings of the same variation, in one more rarely than in the other, and we can imagine that, in the course of time, in these two species, coloured lines over the oblique stripes will arise. In any case these spots are the elements of variation, out of which coloured lines MAY be evolved, if they are combined in this direction through the agency of natural selection. In S. populi the spots are often small, but sometimes it seems as though several had united to form large spots. Whether a process of selection in this direction will arise in S. populi and S. ocellata, or whether it is now going on cannot be determined, since we cannot tell in advance what biological value the marking might have for these two species. It is conceivable that the spots may have no selection- value as far as these species are concerned, and may therefore disappear again in the course of phylogeny, or, on the other hand, that they may be changed in another direction, for instance towards imitation of the rust- red fungoid patches on poplar and willow leaves. In any case we may regard the smallest spots as the initial stages of variation, the larger as a cumulative summation of these. Therefore either these initial stages must already possess selection-value, or, as I said before: THERE MUST BE SOME OTHER REASON FOR THEIR CUMULATIVE SUMMATION. I should like to give one more example, in which we can infer, though we cannot directly observe, the initial stages.
For the present I need only draw the conclusion that one and the same caterpillar may exhibit the initial stages of both, and that it depends on the manner in which these marking elements are INTENSIFIED and COMBINED by natural selection whether whitish longitudinal or oblique stripes should result. In this case then the "useful variations" were actually "always there," and we see that in the same group of Lepidoptera, e.g. species of Sphingidae, evolution has occurred in both directions according to whether the form lived among grass or on broad leaves with oblique lateral veins, and we can observe even now that the species with oblique stripes have longitudinal stripes when young, that is to say, while the stripes have no biological significance. The white places in the skin which gave rise, probably first as small spots, to this protective marking could be combined in one way or another according to the requirements of the species. They must therefore either have possessed selection-value from the first, or, if this was not the case at their earliest occurrence, there must have been SOME OTHER FACTORS which raised them to the point of selection-value. I shall return to this in discussing germinal selection. But the case may be followed still farther, and leads us to the same alternative on a still more secure basis.
Many years ago I observed in caterpillars of Smerinthus populi (the poplar hawk-moth), which also possess white oblique stripes, that certain individuals showed RED SPOTS above these stripes; these spots occurred only on certain segments, and never flowed together to form continuous stripes. In another species (Smerinthus tiliae) similar blood-red spots unite to form a line-like coloured seam in the last stage of larval life, while in S. ocellata rust-red spots appear in individual caterpillars, but more rarely than in S. Populi, and they show no tendency to flow together.
Thus we have here the origin of a new character, arising from small beginnings, at least in S. tiliae, in which species the coloured stripes are a normal specific character. In the other species, S. populi and S. ocellata, we find the beginnings of the same variation, in one more rarely than in the other, and we can imagine that, in the course of time, in these two species, coloured lines over the oblique stripes will arise. In any case these spots are the elements of variation, out of which coloured lines MAY be evolved, if they are combined in this direction through the agency of natural selection. In S. populi the spots are often small, but sometimes it seems as though several had united to form large spots. Whether a process of selection in this direction will arise in S. populi and S. ocellata, or whether it is now going on cannot be determined, since we cannot tell in advance what biological value the marking might have for these two species. It is conceivable that the spots may have no selection- value as far as these species are concerned, and may therefore disappear again in the course of phylogeny, or, on the other hand, that they may be changed in another direction, for instance towards imitation of the rust- red fungoid patches on poplar and willow leaves. In any case we may regard the smallest spots as the initial stages of variation, the larger as a cumulative summation of these. Therefore either these initial stages must already possess selection-value, or, as I said before: THERE MUST BE SOME OTHER REASON FOR THEIR CUMULATIVE SUMMATION. I should like to give one more example, in which we can infer, though we cannot directly observe, the initial stages.