The difficulty of the initial variation had also occurred to Darwin, and he discusses it in an interesting passage which is so important that we may quote it here in full:
It should be observed that the process of imitation probably never commenced between forms widely dissimilar in colour. But starting with species already somewhat like each other, the closest resemblance, if beneficial, could readily be gained by the above means; and if the imitated form was subsequently and gradually modified through any agency, the imitating form would be led along the same track, and thus be altered to almost any extent, so that it might ultimately assume an appearance or colouring wholly unlike that of the other members of the family to which it belonged. There is, however, some difficulty on this head, for it is necessary to suppose in some cases that ancient members belonging to several distinct groups, before they had diverged to their present extent, accidentally resembled a member of another and protected group in a sufficient degree to afford some slight protection; this having given the basis for the subsequent acquisition of the most perfect resemblance[[32]].
Both Darwin and Wallace recognised clearly this difficulty of the initial variations, and both suggested a means of getting over it on similar lines. Both supposed that in general colour and pattern the groups to which model and mimic belonged were far more alike originally than they are to-day. They were in fact so much alike that comparatively small variations in a favourable direction on the part of the mimic would lead to its being confused with the unpalatable model. Then as the model became more and more conspicuously coloured, as it developed a more and more striking pattern warning would-be enemies of its unpleasant taste, the mimic gradually kept pace with it through the operation of natural selection, in the shape of the discriminating enemy, eliminating those most unlike the model. The mimic travelled closely in the wake of the model, coaxed as it were by natural selection, till at last it was far removed in general appearance from the great majority of its near relations.
In this way was offered a comparatively simple method of getting over the difficulty of applying the principle of natural selection to the initial variations in a mimetic approach on the part of one species to another. But it did not escape Darwin's penetration that such an argument would not always be easy of application—that there might be cases where a given model was mimicked by members of several groups of widely differing ancestral pattern, and that in these cases it would be difficult to conceive of members of each of the several groups shewing simultaneous variations which would render them liable to be mistaken for the protected model. The difficulty may perhaps be best illustrated if we consider a definite case.
It is a feature of mimetic resemblances among butterflies that a given species in a given locality may serve as a model for several other species belonging to unrelated groups. Generally such mimics belong to presumably palatable species, but other presumably unpalatable species may also exhibit a similar coloration and pattern. In this way is formed a combine to which the term "mimicry ring" has sometimes been applied. An excellent example of such a mimicry ring is afforded by certain species of butterflies in Ceylon, and is illustrated on [Plate IV]. It is made up in the first place of two species belonging to the presumably distasteful Danaine group, viz. Danais chrysippus and D. plexippus. The latter is a rather darker insect but presents an unmistakable general likeness to D. chrysippus. Those who believe in
Müllerian mimicry would regard it as an excellent example of that phenomenon. For those who believe only in Batesian mimicry D. plexippus, being the scarcer insect, must be regarded as the mimic and D. chrysippus as the model. In both of these species the sexes are similar, whereas in the other three members of the "ring" the female alone exhibits the resemblance. One of these three species is the common Nymphaline, Hypolimnas misippus, of which the female bears an extraordinary likeness to D. chrysippus when set and pinned out on cork in the ordinary way. The male, however ([Pl. IV], fig. 8), is an insect of totally different appearance. The upper surfaces of the wings are velvety black with a large white patch bordered with purple in the middle of each[[33]]. The "ring" is completed by the females of Elymnias undularis and Argynnis hyperbius. The former of these belongs to the group of Satyrine butterflies and the female is usually regarded as a mimic of D. plexippus, which it is not unlike in so far as the upper surface of the wings is concerned. Here again the male is an insect of totally dissimilar appearance. Except for a border of lighter brown along the outer edges of the hind wings the upper surface is of a uniform deep purple-brown
all over ([Pl. IV], fig. 6). In Argynnis hyperbius the appearance is in general that of the Fritillary group to which it belongs. But in the female the outer portion of the fore wings exhibits much black pigment and is crossed by a broad white band similar to that found in the same position on the wing of D. plexippus ([Pl. IV], fig. 2).
Of the five species constituting this little "mimicry ring" in Ceylon two, on the current theory of mimicry, are to be regarded as definitely unpalatable, one (H. misippus) as doubtfully so, while the Satyrine and the Fritillary are evidently examples of simple or Batesian mimicry.
Now such examples as this of simultaneous mimicry in several species are of peculiar interest for us when we come to inquire more closely into the process by which the resemblances can be supposed to have been brought about. Take for example the case of E. undularis. The male is evidently an unprotected insect in so far as mimicry is concerned, while the female exhibits the general pattern and coloration characteristic of the warningly coloured and presumably distasteful species D. plexippus or D. chrysippus. If we are to suppose this to have been brought about by the operation of natural selection it is clear that we must regard the colour and pattern of the male as the original colour and pattern of both sexes. For natural selection cannot be supposed to have operated in causing the male to pass from a protected to an unprotected condition, or even in causing him