Mimicry in Butterflies/Chapter 5
CHAPTER V
SOME CRITICISMS
The facts related in the last two chapters are sufficient to make it clear that these remarkable resemblances between species belonging as a rule to widely different groups constitute a real phenomenon, and as such demand an explanation. One explanation, that in terms of the theory of mimicry, has already been outlined, and we may now turn to consider it in more detail. Some years ago Wallace[1], combating the suggestion that these instances of resemblance might be mere coincidences, laid down five conditions which he stated were applicable to all such cases, and rendered utterly inadequate any explanation other than in terms of natural selection. These five conditions are of historical interest and may also serve as a peg for sundry criticisms in connection with the mimicry theory. They are as follows:
(1) That the imitative species occur in the same area and occupy the very same station as the imitated.
(2) That the imitators are always the more defenceless.
(3) That the imitators are always less numerous in individuals.
(4) That the imitators differ from the bulk of their allies.
(5) That the imitation, however minute, is external and visible only, never extending to internal characters or to such as do not affect the external appearance.
In offering certain criticisms of the mimicry explanation it will be convenient to do so in connection with these five conditions which Wallace regarded as constant for all cases of mimetic resemblance.
(1) That the imitative species occur in the same area and occupy the very same station as the imitated.
This on the whole is generally true. It is well shewn in some of the most striking cases such as those of the Old-World Papilios that mimic Danaines, or of the Dismorphias and their Ithomiine models. In many of these cases the range of neither model nor mimic is a very wide one, yet the mimic is found strictly inside the area inhabited by the model. Papilio agestor, for instance, is only found where Caduga tytia occurs, nor is P. mendax known outside the area frequented by Euploea rhadamanthus. Even more striking in this respect are some of the Ithomiine-Dismorphia resemblances in the New World. The Ithomiine models are as a rule very local though very abundant. Two hundred miles away the predominant Ithomiine often bears quite a distinct pattern, and when this is the case the mimicking Dismorphia is generally changed in the same sense. But though mimic and model may be found together in the same locality, they do not always occupy the same station in the sense that they fly together. According to Seitz[2] the Dismorphias themselves do not fly with the Ithomiines which they mimic. The occurrence of butterflies is largely conditioned by the occurrence of the plants on which the larva feeds, and this is especially true of the female, which, as has already been noticed, is more commonly mimetic than the male. The female of Papilio polytes, for instance, is found flying where are to be found the wild citronaceous plants on which its larva feeds. On the other hand, its so-called models, Papilio hector and P. aristolochiae, are generally in the proximity of the Aristolochias on which their larvae feed. The two plants are not always found together, so that one frequently comes across areas where P. polytes is very abundant while the models are scarce or absent.
Though in the great majority of cases the imitator and the imitated occur in the same locality, this is not always so. The female of the Fritillary Argynnis hyperbius (Pl. IV, fig. 3), for instance, is exceedingly difficult to distinguish from Danais plexippus when flying, although when at rest the difference between the two is sufficiently obvious. Both insects are plentiful in Ceylon but inhabit different stations. The Danaid is a low-country insect, while the Fritillary is not found until several thousand feet up. The two species affect entirely different stations and hardly come into contact with each other. Where one is plentiful the other is not found. It has been suggested that migratory birds may have come into play in such cases. The bird learns in the low country that D. plexippus is unpleasant, and when it pays a visit to the hills it takes this experience with it and avoids those females of the Fritillary which recall the unpleasant Danaine.
Migratory birds have also been appealed to in another case where the resembling species are even further removed from one another than in the last case. Hypolimnas misippus is common and widely spread over Africa and Indo-Malaya, and the male (Pl. IV, fig. 8) bears a simple and conspicuous pattern—a large white spot bordered with purple on each of the very dark fore and hind wings. The same pattern occurs in the males of two other Nymphalines allied to H. misippus, viz. Athyma punctata and Limenitis albomaculata. The two species, however, have a distribution quite distinct from that of H. misippus, being found in China. It has nevertheless been suggested by Professor Poulton[3] that the case may yet be one of mimicry. According to his explanation, H. misippus is unpalatable, the well-known association of its female with Danais chrysippus being an instance of Müllerian mimicry. Migratory birds did the rest. Having had experience of H. misippus in the south, on their arrival in China they spared such specimens of Athyma punctata and Limenitis albomaculata as approached most nearly to H. misippus in pattern, and so brought about the resemblance. The explanation is ingenious, but a simpler view will probably commend itself to most. Other cases are known in which two butterflies bear a close resemblance in pattern and yet are widely separated geographically. Several species of the S. American Vanessid genus Adelpha are in colour scheme like the African Planema poggei which serves as a model for more than one species. The little S. American Phyciodes leucodesma would almost certainly be regarded either as a model for or a mimic of the African Neptis nemetes, did the two occur together. Nevertheless examples of close resemblance between butterflies which live in different parts of the world are relatively rare and serve to emphasise the fact that the great bulk of these resemblance cases are found associated in pairs or in little groups.
(2) That the imitators are always the more defenceless.
In the case of butterflies "defence" as a rule denotes a disagreeable flavour rendering its possessor distasteful to birds and perhaps to other would-be devourers. Feeding experiments with birds (cf. Chapter IX) suggest that certain groups of butterflies, notably the Danaines, Acraeines, Heliconines, Ithomiines and Pharmacophagus Papilios—groups from which models are generally drawn—are characterised by a disagreeable taste, while as a rule this is not true for the mimics. This distasteful quality is frequently accompanied by a more or less conspicuous type of coloration, though this is by no means always so. Many Euploeas are sombre inconspicuous forms, and it is only some of the Ithomiines that sport the gay colours with which that group is generally associated. The members of the distasteful groups usually present certain other peculiarities. Their flight is slower, they are less wary, their bodies are far tougher, and they are more tenacious of life. The slow flight is regarded as an adaptation for exhibiting the warning coloration to the best advantage, but from the point of view of utility it is plausible to suggest that the insect would be better off if in addition to its warning coloration it possessed also the power of swift flight[4]. It is possible that the peculiar slowness of flight of these unpalatable groups is necessitated by the peculiar tough but elastic integument which may present an insufficiently firm and resistant skeletal basis for sharp powerful muscular contraction, and so render swift flight impossible. It is stated that the flight of the mimics is like that of the model, and in some cases this is undoubtedly true. But in a great many cases it certainly does not hold good. Papilio clytia (Pl. I, figs. 7 and 8) is a strong swift flyer very unlike the Danaine and Euploeine which it is supposed to mimic. The flight of the female of Hypolimnas misippus (Pl. IV, fig. 7) is quite distinct from that of Danais chrysippus, while the mimetic forms of P. polytes fly like the non-mimetic one, a mode of flight so different from that of the two models that there is no difficulty in distinguishing them many yards away. Swift flight must be reckoned as one of the chief modes of defence in a butterfly, and on this score the mimic is often better off than the model. And of course it must not be forgotten that where the mode of flight is distinct the protective value of the resemblance must be very much discounted.
(3) That the imitators are always less numerous in individuals.
In the majority of cases this is certainly true. Probably all the Old-World Papilios that mimic Danaines are scarcer, and frequently very much scarcer, than their models. This is very evident from a study of the more comprehensive priced catalogues of Lepidoptera. The mimic is generally a more expensive insect than the model, and not infrequently it costs as many pounds as the model does shillings. But the rule is not universal. Papilio polytes is often much more common than either of its models. The remarkable Pierines, Archonias tereas and A. critias (Pl. XI, fig. 10) as a rule far outnumber the Pharmacophagus Swallow-tail which they mimic. Or again the Chalcosid moth Callamesia pieridoides[5] is a more abundant insect than the Bornean Pierine Delias cathara which it closely resembles.
It has sometimes been suggested in explanation of the greater abundance of the mimic that in such cases we are concerned with Müllerian mimicry, that since both of the species concerned are distasteful there is not, strictly speaking, either a mimic or a model, and consequently the relative proportions have not the significance that they possess where the mimicry is of the simple Batesian type. It is, however, very doubtful whether such an explanation is of any value, for, as will appear later, there are grave objections to accepting the current theory as to the way in which a resemblance is established on Müllerian lines (cf. pp. 72-74).
(4) That the imitators differ from the bulk of their allies.
What importance we attach to this condition must depend upon our interpretation of the word "allies"—whether, for example, we use it for a small group of closely connected species, for a genus, for a group of genera, or in an even wider sense. Perhaps an example will serve to make the difficulty more clear. As already noticed, the S. American genus Dismorphia belongs to the family of Pieridae or "whites." Also certain species of Dismorphia bear a close resemblance to certain species of Ithomiines, a noteworthy example being D. praxinoe and Mechanitis saturata (Pl. X, figs. 3 and 7), in which the pattern, colour, and shape of the two species are all far removed from what is usually understood by a "white." It must not be forgotten, however, that these matters are generally discussed by European naturalists who have grown up in a region where the majority of the "whites" are more or less white. For this reason the statement that D. praxinoe differs from the bulk of its allies is likely to meet with general acceptance, especially as some of the species of the genus itself (e.g., D. cretacea, Pl. X, fig. 1) are regular whites in appearance. But when we come to look at the genus Dismorphia as a whole the matter assumes another complexion. Seitz[6] recognises 75 species of which about a dozen are predominantly white. The rest present a wonderful diversity of colour and pattern. Black predominates on the fore wings, and the insect is frequently marked with gay patches of yellow, bright brown, scarlet, or blue. Forms which from their colour are clearly not mimics present nevertheless the general pattern and shape of other forms which bear a strong resemblance to some Ithomiine. Sometimes a change of colour in certain patches from blue or yellow to bright brown would make all the difference between a non-imitative and an imitative species. Moreover, the non-imitative forms frequently have the peculiar narrow wing, so unusual in a Pierine, which enhances the resemblance of the mimicking species to the Ithomiine model, and which to some extent occurs even in D. cretacea. Clearly we are not justified in saying that D. praxinoe differs from the bulk of its allies, for inside the genus there are many non-imitative species which differ from it in some particulars and are alike it in others. There is a distinct family resemblance among the bulk of the Dismorphias, including practically all the mimetic forms, and on the whole the resemblances between the imitative and the non-imitative forms are as noteworthy as the differences. Though not exhibited in so striking a fashion, the same is to a large extent true of a large proportion of the cases of mimicry. It is on the whole unusual to find cases where a single species departs widely from the pattern scheme of the other members of the genus and at the same time resembles an unrelated species. Two of the best instances are perhaps those of Limenitis archippus (p. 49) and of the Pierid Pareronia (p. 23). Of the total number of mimicry instances a high proportion is supplied by relatively few groups. In each region several main series of models and mimics run as it were parallel to one another. In Asia, for example, we have the Papilio-Danaine series where the colour-patterns of a series of Danaines, all nearly related, are closely paralleled by those of a section of the genus Papilio, and by those of the Satyrid genus Elymnias. In Africa there is a similar Papilio-Danaine series though of less extent. Africa has a group of models not found in Asia, and the Papilio-Danaine series is as it were curtailed by the Papilio-Planema series with which to some extent runs parallel the genus Pseudacraea. These phenomena of parallel series have been mentioned here as shewing that mimicry tends to run in certain groups and that in many cases at any rate little meaning can be attached to the statement that the imitators differ from the bulk of their allies.
The fifth of Wallace's conditions is clear and needs no discussion.
It is evident that at any rate a large proportion of the instances of close resemblance do not fulfil all of the conditions laid down by Wallace. Nevertheless we should expect them to do so if the resemblance has been brought about by the cumulative effect of natural selection on small favourable variations. Clearly there is a prima facie case for doubting whether we must of necessity ascribe all resemblance of the kind to natural selection, and in the next few chapters we shall discuss it in more detail from several points of view.
- ↑ Darwinism, 1890 (1st Edition 1889), p. 264.
- ↑ Macrolepidoptera of the World. Fauna Americana, p. 98.
- ↑ Essays on Evolution, 1908, p. 381.
- ↑ These "unpalatable" butterflies are sometimes extensively preyed upon by insectivorous birds, when they fall an easier prey owing to their slowness (cf. p. 112).
- ↑ See Shelford, Proc. Zool. Soc. 1902, p. 260. A coloured figure of both species is given in the paper.
- ↑ Macrolepidoptera of the World. Fauna Americana, pp. 98-104, Plates 28-30.