While staying a few days, in July 1887, at the Summit Hotel on the Central Pacific Railway, I strolled out one evening after dinner, and on the road, not fifty yards from the house, I saw a pretty little white and black animal with a bushy tail coming towards me. As it came on at a slow pace and without any fear, although it evidently saw me, I thought at first that it must be some tame creature, when it suddenly occurred to me that it was a skunk. It came on till within five or six yards of me, then quietly climbed over a dwarf wall and disappeared under a small outhouse, in search of chickens, as the landlord afterwards told me. This animal possesses, as is well known, a most offensive secretion, which it has the power of ejecting over its enemies, and which effectually protects it from attack. The odour of this substance is so penetrating that it taints, and renders useless, everything it touches, or in its vicinity. Provisions near it become uneatable, and clothes saturated with it will retain the smell for several weeks, even though they are repeatedly washed and dried. A drop of the liquid in the eyes will cause blindness, and Indians are said not unfrequently to lose their sight from this cause. Owing to this remarkable power of offence the skunk is rarely attacked by other animals, and its black and white fur, and the bushy white tail carried erect when disturbed, form the danger-signals by which it is easily distinguished in the twilight or moonlight from unprotected animals. Its consciousness that it needs only to be seen to be avoided gives it that slowness of motion and fearlessness of aspect which are, as we shall see, characteristic of most creatures so protected.
Warning Colours among Insects.
It is among insects that warning colours are best developed, and most abundant. We all know how well marked and conspicuous are the colours and forms of the stinging wasps and bees, no one of which in any part of the world is known to be protectively coloured like the majority of defenceless insects. Most of the great tribe of Malacoderms among beetles are distasteful to insect-eating animals. Our red and black Telephoridae, commonly called "soldiers and sailors," were found, by Mr. Jenner Weir, to be refused by small birds. These and the allied Lampyridae (the fireflies and glow-worms) in Nicaragua, were rejected by Mr. Belt's tame monkey and by his fowls, though most other insects were greedily eaten by them. The Coccinellidae or lady-birds are another uneatable group, and their conspicuous and singularly spotted bodies serve to distinguish them at a glance from all other beetles.
These uneatable insects are probably more numerous than is supposed, although we already know immense numbers that are so protected. The most remarkable are the three families of butterflies—Heliconidae, Danaidae, and Acraeidae—comprising more than a thousand species, and characteristic respectively of the three great tropical regions—South America, Southern Asia, and Africa. All these butterflies have peculiarities which serve to distinguish them from every other group in their respective regions. They all have ample but rather weak wings, and fly slowly; they are always very abundant; and they all have conspicuous colours or markings, so distinct from those of other families that, in conjunction with their peculiar outline and mode of flight, they can usually be recognised at a glance. Other distinctive features are, that their colours are always nearly the same on the under surface of their wings as on the upper; they never try to conceal themselves, but rest on the upper surfaces of leaves or flowers; and, lastly, they all have juices which exhale a powerful scent, so that when one kills them by pinching the body, the liquid that exudes stains the fingers yellow, and leaves an odour that can only be removed by repeated washings.
Now, there is much direct evidence to show that this odour, though not very offensive to us, is so to most insect-eating creatures. Mr. Bates observed that, when set out to dry, specimens of Heliconidae were less subject to the attacks of vermin; while both he and I noticed that they were not attacked by insect-eating birds or dragonflies, and that their wings were not found in the forest paths among the numerous wings of other butterflies whose bodies had been devoured. Mr. Belt once observed a pair of birds capturing insects for their young; and although the Heliconidae swarmed in the vicinity, and from their slow flight could have been easily caught, not one was ever pursued, although other butterflies did not escape. His tame monkey also, which would greedily munch up other butterflies, would never eat the Heliconidae. It would sometimes smell them, but always rolled them up in its hand and then dropped them.
We have also some corresponding evidence as to the distastefulness of the Eastern Danaidae. The Hon. Mr. Justice Newton, who assiduously collected and took notes upon the Lepidoptera of Bombay, informed Mr. Butler of the British Museum that the large and swift-flying butterfly Charaxes psaphon, was continually persecuted by the bulbul, so that he rarely caught a specimen of this species which had not a piece snipped out of the hind wings. He offered one to a bulbul which he had in a cage, and it was greedily devoured, whilst it was only by repeated persecution that he succeeded in inducing the bird to touch a Danais.[92]
Besides these three families of butterflies, there are certain groups of the great genus Papilio—the true swallow-tailed butterflies—which have all the characteristics of uneatable insects. They have a special coloration, usually red and black (at least in the females), they fly slowly, they are very abundant, and they possess a peculiar odour somewhat like that of the Heliconidae. One of these groups is common in tropical America, another in tropical Asia, and it is curious that, although not very closely allied, they have each the same red and black colours, and are very distinct from all the other butterflies of their respective countries. There is reason to believe also that many of the brilliantly coloured and weak-flying diurnal moths, like the fine tropical Agaristidae and burnet-moths, are similarly protected, and that their conspicuous colours serve as a warning of inedibility. The common burnet-moth (Anthrocera filipendula) and the equally conspicuous ragwort-moth (Euchelia jacobeae) have been proved to be distasteful to insect-eating creatures.
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 larvae 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.[93] 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 caterpillars—Abraxas grossulariata, Diloba caeruleocephala, Anthrocera filipendula, and Cucullia verbasci. He also found that they would not touch any hairy or spiny larvae, 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.[94]