The whole of the present investigation had not been planned when I joined with those who, from the first, admitted the omnipotence of natural selection as an article of faith or scientific axiom. A question which can only be solved by the inductive method cannot possibly be regarded as settled, nor can further evidence be considered unnecessary, because the first proofs favour the principle. The admission of a mysteriously working phyletic power appears very unsatisfactory to those who are striving after knowledge; the existence of this power, however, is not to be considered as disproved, because hundreds of characters can be referred to the action of natural selection, and many others to that of the direct action of the conditions of life. If the development of the organic world is to be considered as absolutely dependent on the influence of the environment, not only should we be able here and there to select at pleasure characters which appeared the most accessible for elucidating this point, but it becomes in the first place necessary to attempt to completely refer all characters belonging to any particular group of phenomena, however small this group might be, to known transforming factors. We should then see whether this were possible, or whether there would remain residual phenomena not explicable by known principles and compelling us to admit the existence of a force of development innate in the organism. In any case the “phyletic vital force” can only be got rid of by a process of elimination—by proving that all the characters generally occurring throughout the group of phenomena in question, must be attributed to other causes, and that consequently nothing remains for the action of the supposed phyletic vital force, which would in this manner be negatived, since we cannot infer the presence of a force if the latter exerts no action whatever.
I shall here attempt such an investigation of the group of phenomena displayed by larval markings, with special reference to those of the Sphingidæ. The alternatives upon which we have to decide are the following:—Are the markings of caterpillars purely morphological characters, produced entirely by internal causes? or, are they simply the response of the organism to external influences?
The solution of these questions will be arrived at by seeking to refer all the markings present to one of the known transforming factors, and the success or failure of this attempt will give the required decision. The first question to be attacked is obviously this,—whether the Sphinx-markings are actually, as they appear at first sight, purely morphological characters. If it can be shown that all these markings were originally of biological value, they must be attributed to the action of natural selection.
Did I here at once proceed to establish the biological value of larval markings—and especially of those of the Sphingidæ—so as to arrive in this manner at a conclusion as to their dependence upon natural selection, it would be impossible to avoid the consideration of the total coloration of the caterpillars, since the marking frequently consists only of a local strengthening of the colour, and cannot be comprehended without coming to this understanding. The action of the markings also often appears to be opposed to that of the colouring, making the caterpillar again conspicuous; so that the two factors must necessarily be considered together. I shall therefore commence the investigation with colour in general, and then proceed to treat of marking.
IV.
Biological Value of Colour.
The general prevalence of protective colouring among caterpillars has already been so frequently treated of that it is not here my intention to recall particular instances. In order to judge of the effect of marking, however, it will be well to bear in mind that these insects, being generally defenceless and thus requiring protection, have acquired the most diverse means of rendering themselves in some measure secure from their foes.
The sharp spines which occur on the caterpillars of many butterflies (Vanessa, Melitæa, Argynnis), and the hairs on those of many moths, serve for protective purposes. Among other means of protection—although in a different sense—we have in all the species of the great family of the Papilionidæ the strikingly coloured (yellowish red) odour-emitting tentacles concealed near the head, and suddenly protruded for terrifying foes; and likewise the forked horn at the tail of the caterpillars of the genus of moths Harpyia, the tentacles of which can be suddenly protruded in a similar manner. Adaptive colours and forms combined with certain habits[129] are, however, much more common than defensive weapons. Thus, the caterpillars of the Noctuæ belonging to the genus Catocala and its allies, feed only at night on the green leaves of various forest-trees; by day they rest in crevices of the bark on the tree trunk, which they resemble so perfectly in the colour of their peculiar glossy dull grey or brownish skin beset with small humps, that only sharp eyes can detect them, even when we are familiar with their habits.[130]
The striking resemblance of many moths to splinters of wood is well known, and to this is added a habit which helps their disguise, viz., that of remaining stiff and motionless on the approach of danger, just like a splinter projecting from the branch.[131] Among the moths coming under this category may be mentioned Cucullia Verbasci, and particularly those of the genus Xylina, which, when at rest, closely resemble a broken splinter of wood in the colour and marking of their fore wings, and when touched, have a habit of drawing in their legs and falling without opening their wings as though dead.