The most interesting and most conclusive example of warning coloration is, however, furnished by caterpillars, because in this case the facts have been carefully ascertained experimentally by competent observers. In the year 1866, when Mr. Darwin was collecting evidence as to the supposed effect of sexual selection in bringing about the brilliant coloration of the higher animals, he was struck by the fact that many caterpillars have brilliant and conspicuous colours, in the production of which sexual selection could have no place. We have numbers of such caterpillars in this country, and they are characterised not only by their gay colours but by not concealing themselves. Such are the mullein and the gooseberry caterpillars, the larvæ of the spurge hawk-moth, of the buff-tip, and many others. Some of these caterpillars are wonderfully conspicuous, as in the case of that noticed by Mr. Bates in South America, which was four inches long, banded across with black and yellow, and with bright red head, legs, and tail. Hence it caught the eye of any one who passed by, even at the distance of many yards.

Mr. Darwin asked me to try and suggest some explanation of this coloration; and, having been recently interested in the question of the warning coloration of butterflies, I suggested that this was probably a similar case,-that these conspicuous caterpillars were distasteful to birds and other insect-eating creatures, and that their bright non-protective colours and habit of exposing themselves to view, enabled their enemies to distinguish them at a glance from the edible kinds and thus learn not to touch them; for it must be remembered that the bodies of caterpillars while growing are so delicate, that a wound from a bird's beak would be perhaps as fatal as if they were devoured.1 At this time not a single experiment or observation had been made on the subject, but after I had brought the matter before the Entomological Society, two gentlemen, who kept birds and other tame animals, undertook to make experiments with a variety of caterpillars.

Mr. Jenner Weir was the first to experiment with ten species of small birds in his aviary, and he found that none of them would eat the following smooth-skinned conspicuous cater

1 See Darwin's Descent of Man, p. 325.

pillars-Abraxas grossulariata, Diloba cæruleocephala, Anthrocera filipendula, and Cucullia verbasci. He also found that they would not touch any hairy or spiny larvæ, and he was satisfied that it was not the hairs or the spines, but the unpleasant taste that caused them to be rejected, because in one case a young smooth larva of a hairy species, and in another case the pupa of a spiny larva, were equally rejected. On the other hand, all green or brown caterpillars as well as those that resemble twigs were greedily devoured.1

Mr. A. G. Butler also made experiments with some green lizards (Lacerta viridis), which greedily ate all kinds of food, including flies of many kinds, spiders, bees, butterflies, and green caterpillars; but they would not touch the caterpillar of the gooseberry-moth (Abraxas grossulariata), or the imago of the burnet-moth (Anthrocera filipendula). The same thing happened with frogs. When the gooseberry caterpillars were first given to them, "they sprang forward and licked them eagerly into their mouths; no sooner, however, had they done so, than they seemed to become aware of the mistake that they had made, and sat with gaping mouths, rolling their tongues about, until they had got quit of the nauseous morsels, which seemed perfectly uninjured, and walked off as briskly as ever." Spiders seemed equally to dislike them. This and another conspicuous caterpillar (Halia wavaria) were rejected by two species-the geometrical garden spider (Epeira diadema) and a hunting spider.2

Some further experiments with lizards were made by Professor Weismann, quite confirming the previous observations; and in 1886 Mr. E. B. Poulton of Oxford undertook a considerable series of experiments, with many other species of larvæ and fresh kinds of lizards and frogs. Mr. Poulton then reviewed the whole subject, incorporating all recorded facts, as well as some additional observations made by Mr. Jenner Weir in 1886. More than a hundred species of larvæ or of perfect insects of various orders have now been made the subject of experiment, and the results completely confirm my original suggestion. In almost every case the protectively coloured larvæ have been greedily eaten by all kinds of insectivorous 1 Transactions of the Entomological Society of London, 1869, p. 21. 2 Ibid., p. 27.

animals, while, in the immense majority of cases, the conspicuous, hairy, or brightly coloured larvæ have been rejected by some or all of them. In some instances the inedibility of the larvæ extends to the perfect insect, but not in others. In the former cases the perfect insect is usually adorned with conspicuous colours, as the burnet and ragwort moths; but in the case of the buff-tip, the moth resembles a broken piece of rotten stick, yet it is partly inedible, being refused by lizards. It is, however, very doubtful whether these are its chief enemies, and its protective form and colour may be needed against insectivorous birds or mammals.

Mr. Samuel H. Scudder, who has largely bred North American butterflies, has found so many of the eggs and larvæ destroyed by hymenopterous and dipterous parasites that he thinks at least nine-tenths, perhaps a greater proportion, never reach maturity. Yet he has never found any evidence that such parasites attack either the egg or the larva of the inedible Danais archippus, so that in this case the insect is distasteful to its most dangerous foes in all the stages of its existence, a fact which serves to explain its great abundance and its extension over almost the whole world.1

One case has been found of a protectively coloured larva, -one, moreover, which in all its habits shows that it trusts to concealment to escape its enemies-which was yet always rejected by lizards after they had seized it, evidently under the impression that from its colour it would be eatable. This is the caterpillar of the very common moth Mania typica; and Mr. Poulton thinks that, in this case, the unpleasant taste is an incidental result of some physiological processes in the organism, and is itself a merely useless character. It is evident that the insect would not conceal itself so carefully as it does if it had not some enemies, and these are probably birds or small mammals, as its food-plants are said to be dock and willow-herb, not suggestive of places frequented by lizards; and it has been found by experiment that lizards and birds have not always the same likes and dislikes. The case is interesting, because it shows that nauseous fluids sometimes occur sporadically, and may thus be intensified by natural selection when required for the purpose

1 Nature, vol. iii. p. 147. One parasite, Pteromalus archippi, has since been found.

of protection. Another exceptional case is that of the very conspicuous caterpillar of the spurge hawk-moth (Deilephila euphorbia), which was at once eaten by a lizard, although, as it exposes itself on its food-plant in the daytime and is very abundant in some localities, it must almost certainly be disliked by birds or by some animals who would otherwise devour it. If disturbed while feeding it is said to turn round with fury and eject a quantity of green liquid, of an acid and disagreeable smell similar to that of the spurge milk, only worse.1

These facts, and Mr. Poulton's evidence that some larvæ rejected by lizards at first will be eaten if the lizards are very hungry, show that there are differences in the amount of the distastefulness, and render it probable that if other food were wanting many of these conspicuous insects would be eaten. It is the abundance of the eatable kinds that gives value to the inedibility of the smaller number; and this is probably the reason why so many insects rely on protective colouring rather than on the acquisition of any kind of defensive weapons. In the long run the powers of attack and defence must balance each other. Hence we see that even the powerful stings of bees and wasps only protect them against some enemies, since a tribe of birds, the bee-eaters, have been developed which feed upon them, and some frogs and lizards do so occasionally

The preceding outline will sufficiently explain the characteristics of "warning coloration" and the end it serves in nature. There are many other curious modifications of it, but these will be best appreciated after we have discussed the remarkable phenomenon of "mimicry," which is bound up with and altogether depends upon "warning colour," and is in some cases the chief indication we have of the possession of some offensive weapon to secure the safety of the species imitated.

Mimicry.

This term has been given to a form of protective resemblance, in which one species so closely resembles another in external form and colouring as to be mistaken for it, although the two may not be really allied and often belong to distinct

1 Stainton's Manual of Butterflies and Moths, vol. i. p. 93; E. B. Poulton, Proceedings of the Zool. Soc. of London, 1887, pp. 191-274.

families or orders. One creature seems disguised in order to be made like another; hence the terms "mimic" and mimicry, which imply no voluntary action on the part of the imitator. It has long been known that such resemblances do occur, as, for example, the clear-winged moths of the families Sesiidae and Ægeriidæ, many of which resemble bees, wasps, ichneumons, or saw-flies, and have received names expressive of the resemblance; and the parasitic flies (Volucella) which closely resemble bees, on whose larvæ the larvæ of the flies feed.

The great bulk of such cases remained, however, unnoticed, and the subject was looked upon as one of the inexplicable curiosities of nature, till Mr. Bates studied the phenomenon among the butterflies of the Amazon, and, on his return home, gave the first rational explanation of it.1 The facts are, briefly, these. Everywhere in that fertile region for the entomologist the brilliantly coloured Heliconidæ abound, with all the characteristics which I have already referred to when describing them as illustrative of "warning coloration." But along with them other butterflies were occasionally captured, which, though often mistaken for them, on account of their close resemblance in form, colour, and mode of flight, were found on examination to belong to a very distinct family, the Pierida. Mr. Bates notices fifteen distinct species of Pieridæ, belonging to the genera Leptalis and Euterpe, each of which closely imitates some one species of Heliconidæ, inhabiting the same region and frequenting the same localities. It must be remembered that the two families are altogether distinct in structure. The larvæ of the Heliconidæ are tubercled or spined, the pupæ suspended head downwards, and the imago has imperfect forelegs in the male; while the larvæ of the Pieridæ are smooth, the pupæ are suspended with a brace to keep the head erect, and the forefeet are fully developed in both sexes. These differences are as large and as important as those between pigs and sheep, or between swallows and sparrows; while English entomologists will best understand the case by supposing that a species of Pieris in this country was coloured and shaped like a small tortoise-shell, while another species on the Continent was equally like a Camberwell beauty-so like in both

1 See Transactions of the Linnean Society, vol. xxiii. pp. 495-566, coloured plates.