Mimicry in Butterflies/Chapter 2
CHAPTER II
MIMICRY—BATESIAN AND MÜLLERIAN
Mimicry is a special branch of the study of adaptation. The term has sometimes been used loosely to include cases where an animal, most frequently an insect, bears a strong and often most remarkable resemblance to some feature of its inanimate surroundings. Many butterflies with wings closed are wonderfully like dead leaves; certain spiders when at rest on a leaf look exactly like bird-droppings; "looper" caterpillars simulate small twigs; the names of the "stick-" and "leaf-" insects are in themselves an indication of their appearance. Such cases as these, in which the creature exhibits a resemblance to some part of its natural surroundings, should be classified as cases of "protective resemblance" in contradistinction to mimicry proper. Striking examples of protective resemblance are abundant, and though we possess little critical knowledge of the acuity of perception in birds and other insect feeders it is plausible to regard the resemblances as being of definite advantage in the struggle for existence. However, it is with mimicry and not with protective coloration in general that we are here directly concerned, and the nature of the phenomenon may perhaps best be made clear by a brief account of the facts which led to the statement of the theory.
In the middle of last century the distinguished naturalist, H. W. Bates, was engaged in making collections in parts of the Amazon region. He paid much attention to butterflies, in which group he discovered a remarkably interesting phenomenon[1]. Among the species which he took were a large number belonging to the group Ithomiinae, small butterflies of peculiar appearance with long slender bodies and narrow wings bearing in most cases a conspicuous pattern (cf. Pl. X, fig. 7). When Bates came to examine his catch more closely he discovered that among the many Ithomiines were a few specimens very like them in general shape, colour, and markings, but differing in certain anatomical features by which the Pierinae, or "whites," are separated from other groups. Most Pierines are very different from Ithomiines. It is the group to which our common cabbage butterfly belongs and the ground colour is generally white. The shape of the body and also of the wings is in general quite distinct from what it is in the Ithomiines. Nevertheless in these particular districts certain of the species of Pierines had departed widely from what is usually regarded as their ancestral pattern (Pl. X, fig. 1) and had come to resemble very closely the far more abundant Ithomiines among whom they habitually flew (cf. Pl. X, figs. 2 and 3). To use Bates' term they "mimicked" the Ithomiines, and he set to work to devise an explanation of how this could have come about. The Origin of Species had just appeared and it was natural that Bates should seek to interpret this peculiar phenomenon on the lines there laid down. How was it that these Pierines had come to depart so widely from the general form of the great bulk of their relations, and to mimic so closely in appearance species belonging to an entirely different group, while at the same time conserving the more deeply seated anatomical features of their own family? If the change was to be regarded as having come about through the agency of natural selection it must clearly be of advantage to the mimicking forms; otherwise natural selection could not come into operation. What advantage then have the Ithomiines over the majority of butterflies in those parts? They are small insects, rather flimsy in build, with comparatively weak powers of flight, and yet so conspicuously coloured that they can hardly be mistaken for anything else. In spite of all this they are little subject to the attacks of enemies such as birds, and Bates attributed this to the fact that the juices of their bodies are unpalatable. According to him their striking and conspicuous pattern is of the nature of a warning coloration, advertising their disagreeable properties to possible enemies. A bird which had once attempted to eat one would find it little to its taste. It would thenceforward associate the conspicuous pattern with a disagreeable flavour and in future leave such butterflies severely alone. The more conspicuous the pattern the more readily would it be noticed by the enemy, and so it would be of advantage to the Ithomiine to possess as striking a pattern as possible. Those butterflies shewing a tendency to a more conspicuous pattern would be more immune to the attacks of birds and so would have a better chance of leaving progeny than those with a less conspicuous pattern. In this way variations in the direction of greater conspicuousness would be accumulated gradually by natural selection, and so would be built up in the Ithomiine the striking warning coloration by which it advertises its disagreeable properties. Such is the first step in the making of a mimicry case—the building up through natural selection of a conspicuous pattern in an unpalatable species by means of which it is enabled to advertise its disagreeable properties effectively and thereby secure immunity from the attacks of enemies which are able to appreciate the advertisement. Such patterns and colours are said to be of a "warning" nature. The existence of an unpalatable model in considerable numbers is the first step in the production of a mimetic resemblance through the agency of natural selection.
We come back now to our Pierine which must be assumed to shew the general characters and coloration of the family of whites to which they belong (cf. Pl. X, fig. 1). Theoretically they are not specially protected by nauseous properties from enemies and hence their conspicuous white coloration renders them especially liable to attack. If, however, they could exchange their normal dress for one resembling that of the Ithomiines it is clear that they would have a chance of being mistaken for the latter and consequently of being left alone. Moreover, in certain cases these Pierines have managed to discard their normal dress and assume that of the Ithomiines. On theoretical grounds this must clearly be of advantage to them, and being so might conceivably have arisen through the operation of natural selection. This indeed is what is supposed to have taken place on the theory of mimicry. Those Pierines which exhibited a variation of colour in the direction of the Ithomiine "model" excited distrust in the minds of would-be devourers, who had learned from experience to associate that particular type of coloration with a disagreeable taste. Such Pierines would therefore have a rather better chance of surviving and of leaving offspring. Some of the offspring would exhibit the variation in a more marked degree and these again would in consequence have a yet better chance of surviving. Natural selection would encourage those varying in the direction of the Ithomiine model at the expense of the rest and by its continuous operation there would gradually be built up those beautiful cases of resemblance which have excited the admiration of naturalists.
Wallace was the next after Bates to interest himself in mimicry and, from his study of the butterflies of the Oriental region[2], shewed that in this part of the world too there existed these remarkable resemblances between species belonging to different families. Perhaps the most important part of Wallace's contribution was the demonstration that in some species not only was it the female alone that "mimicked" but that there might be several different forms of female mimicking different models, and in some cases all unlike the male of their own species. One of the species studied by Wallace, Papilio polytes, is shewn on Plate V. We shall have occasion to refer to this case later on, and it is sufficient here to call attention to the three different forms of female, of which one is like the male while the other two resemble two other species of Papilio, P. hector and P. aristolochiae, which occur in the same localities. Instances where the female alone of some unprotected species mimics a model with obnoxious properties are common in all tropical countries. It has been suggested that this state of things has come about owing to the greater need of protection on the part of the female. Hampered by the disposal of the next generation the less protected female would be at a greater disadvantage as compared with the mimic than would the corresponding male whose obligations to posterity are more rapidly discharged. The view of course makes the assumption that the female transmits her peculiar properties to her daughters but not to her sons.
A few years later Trimen[3] did for Africa what Bates had done for America and Wallace for Indo-Malaya. It was in this paper that he elucidated that most remarkable of all cases of mimicry—Papilio dardanus with his harem of different consorts, all tailless, all unlike himself, and often wonderfully similar to unpalatable forms found in the same localities (cf. p. 30).
We may now turn to one of the most ingenious developments of the theory of mimicry. Not long after Bates' original memoir appeared attention was directed to a group of cases which could not be explained on the simple hypothesis there put forward. Many striking cases of resemblance had been adduced in which both species obviously belonged to the presumably unpalatable groups. Instances of the sort had been recorded by Bates himself and are perhaps most plentiful in South America between species belonging respectively to the Ithomiinae and Heliconinae. On the theory of mimicry all the members of both of these groups must be regarded as specially protected owing to their conspicuous coloration and distasteful properties. What advantage then can an Ithomiine be supposed to gain by mimicking a Heliconine, or vice versâ? Why should a species exchange its own bright and conspicuous warning pattern for one which is neither brighter nor more conspicuous? To Fritz Müller, the well-known correspondent of Darwin, belongs the credit of having suggested a way out of the difficulty. Müller's explanation turns upon the education of birds. Every year there hatch into the world fresh generations of young birds, and each generation has to learn afresh from experience what is pleasant to eat and what is not. They will try all things and hold fast to that which is good. They will learn to associate the gay colours of the Heliconine and the Ithomiine with an evil taste[4] and they will thenceforward avoid butterflies which advertise themselves by means of these particular colour combinations. But in a locality where there are many models, each with a different pattern and colour complex, each will have to be tested separately before the unpalatableness of each is realised. If for example a thousand young birds started their education on a population of butterflies in which there were five disagreeable species, each with a distinct warning pattern, it is clear that one thousand of each would devote their lives to the education of these birds, or five thousand butterflies in all[5]. But if these five species, instead of shewing five distinct warning patterns, all displayed the same one it is evident that the education of the birds would be accomplished at the price of but one thousand butterfly existences instead of five. Even if one of the five species were far more abundant than the others it would yet be to its advantage that the other four should exhibit the same warning pattern. Even though the losses were distributed pro rata the more abundant species would profit to some extent. For the less abundant species the gain would of course be relatively greater. Theoretically therefore, all of the five species would profit if in place of five distinct warning patterns they exhibited but a single one in common. And since it is profitable to all concerned what more natural than that it should be brought about by natural selection?
Müller's views are now widely accepted by students of mimicry as an explanation of these curious cases where two or more evidently distasteful species closely resemble one another. Indeed the tendency in recent years has been to see Müllerian mimicry everywhere, and many of the instances which were long regarded as simple Batesian cases have now been relegated to this category. The hypothesis is, of course, based upon what appears to man to be the natural behaviour of young birds under certain conditions. No one knows whether young birds actually do behave in the way that they are supposed to. In the absence of any such body of facts the Müllerian hypothesis cannot rank as more than a plausible suggestion, and, as will appear later, it is open to severe criticism on general grounds.
Perhaps the next contribution to the subject of mimicry which must rank of the first importance was that of Erich Haase[6], to whose book students of these matters must always be under a heavy obligation. It was the first and still remains the chief work of general scope. Since Haase's day great numbers of fresh instances of mimetic resemblance have been recorded from all the great tropical areas of the world, and the list is being added to continually. Most active in this direction is the Oxford School under Professor Poulton to whose untiring efforts are largely due the substantial increases in our knowledge of African butterflies contributed by various workers in the field during the past few years. Whatever the interpretation put upon them, there can be no question as to the value of the facts brought together, more especially those referring to the nature of the families raised in captivity from various mimetic forms. With the considerable additions from Africa[7] during the past few years several hundreds of cases of mimicry must now have been recorded. Some of the best known and most striking from among these will be described briefly in the next two chapters.
- ↑ Trans. Linn. Soc. vol. 23, 1862.
- ↑ Trans. Linn. Soc. vol. 25, 1866.
- ↑ Trans. Linn. Soc. vol. 26, 1870.
- ↑ In attributing this quality to the butterflies in question I am merely stating what is held by the supporters of the mimicry theory. I know of scarcely any evidence either for or against the supposition.
- ↑ It is assumed that the intelligence of the birds is such that they can learn a pattern after a single disagreeable experience of it.
- ↑ Untersuchungen über die Mimikry, 1893.
- ↑ The African mimetic butterflies have been recently monographed by Eltringham in a large and beautifully illustrated work—African Mimetic Butterflies, Oxford, 1910.