Darwin and Modern Science [42]
It is true that this explanation of the bright, conspicuous colours is only a hypothesis, but its foundations,--unpalatableness, and the liability of other butterflies to be eaten,--are certain, and its consequences--the existence of mimetic palatable forms--confirm it in the most convincing manner. Of the many cases now known I select one, which is especially remarkable, and which has been thoroughly investigated, Papilio dardanus (merope), a large, beautiful, diurnal butterfly which ranges from Abyssinia throughout the whole of Africa to the south coast of Cape Colony.
The males of this form are everywhere ALMOST the same in colour and in form of wings, save for a few variations in the sparse black markings on the pale yellow ground. But the females occur in several quite different forms and colourings, and one of these only, the Abyssinian form, is like the male, while the other three or four are MIMETIC, that is to say, they copy a butterfly of quite a different family the Danaids, which are among the IMMUNE forms. In each region the females have thus copied two or three different immune species. There is much that is interesting to be said in regard to these species, but it would be out of keeping with the general tenor of this paper to give details of this very complicated case of polymorphism in P. dardanus. Anyone who is interested in the matter will find a full and exact statement of the case in as far as we know it, in Poulton's "Essays on Evolution" (pages 373-375). (Professor Poulton has corrected some wrong descriptions which I had unfortunately overlooked in the Plates of my book "Vortrage uber Descendenztheorie", and which refer to Papilio dardanus (merope). These mistakes are of no importance as far as and understanding of the mimicry-theory is concerned, but I hope shortly to be able to correct them in a later edition.) I need only add that three different mimetic female forms have been reared from the eggs of a single female in South Africa. The resemblance of these forms to their immune models goes so far that even the details of the LOCAL forms of the models are copied by the mimetic species.
It remains to be said that in Madagascar a butterfly, Papilio meriones, occurs, of which both sexes are very similar in form and markings to the non-mimetic male of P. dardanus, so that it probably represents the ancestor of this latter species.
In face of such facts as these every attempt at another explanation must fail. Similarly all the other details of the case fulfil the preliminary postulates of selection, and leave no room for any other interpretation. That the males do not take on the protective colouring is easily explained, because they are in general more numerous, and the females are more important for the preservation of the species, and must also live longer in order to deposit their eggs. We find the same state of things in many other species, and in one case (Elymnias undularis) in which the male is also mimetically coloured, it copies quite a differently coloured immune species from the model followed by the female. This is quite intelligible when we consider that if there were TOO MANY false immune types, the birds would soon discover that there were palatable individuals among those with unpalatable warning colours. Hence the imitation of different immune species by Papilio dardanus!
I regret that lack of space prevents my bringing forward more examples of mimicry and discussing them fully. But from the case of Papilio dardanus alone there is much to be learnt which is of the highest importance for our understanding of transformations. It shows us chiefly what I once called, somewhat strongly perhaps, THE OMNIPOTENCE OF NATURAL SELECTION in answer to an opponent who had spoken of its "inadequacy." We here see that one and the same species is capable of producing four or five different patterns of colouring and marking; thus the colouring and marking are not, as has often been supposed, a necessary outcome of the specific nature of the species, but a true adaptation,