Contributions to the Theory of Natural Selection. Alfred Russel Wallace

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varied means by which animals are defended or concealed. One of the uses of the phosphorescence with which many insects are furnished, is probably to frighten away their enemies; for Kirby and Spence state that a ground beetle (Carabus) has been observed running round and round a luminous centipede as if afraid to attack it. An immense number of insects have stings, and some stingless ants of the genus Polyrachis are armed with strong and sharp spines on the back, which must render them unpalatable to many of the smaller insectivorous birds. Many beetles of the family Curculionidæ have the wing cases and other external parts so excessively hard, that they cannot be pinned without first drilling a hole to receive the pin, and it is probable that all such find a protection in this excessive hardness. Great numbers of insects hide themselves among the petals of flowers, or in the cracks of bark and timber; and finally, extensive groups and even whole orders have a more or less powerful and disgusting smell and taste, which they either possess permanently, or can emit at pleasure. The attitudes of some insects may also protect them, as the habit of turning up the tail by the harmless rove-beetles (Staphylindidæ) no doubt leads other animals besides children to the belief that they can sting. The curious attitude assumed by sphinx caterpillars is probably a safeguard, as well as the blood-red tentacles which can suddenly be thrown out from the neck, by the caterpillars of all the true swallow-tailed butterflies.

      It is among the groups that possess some of these varied kinds of protection in a high degree, that we find the greatest amount of conspicuous colour, or at least the most complete absence of protective imitation. The stinging Hymenoptera, wasps, bees, and hornets, are, as a rule, very showy and brilliant insects, and there is not a single instance recorded in which any one of them is coloured so as to resemble a vegetable or inanimate substance. The Chrysididæ, or golden wasps, which do not sting, possess as a substitute the power of rolling themselves up into a ball, which is almost as hard and polished as if really made of metal,—and they are all adorned with the most gorgeous colours. The whole order Hemiptera (comprising the bugs) emit a powerful odour, and they present a very large proportion of gay-coloured and conspicuous insects. The lady-birds (Coccinellidæ) and their allies the Eumorphidæ, are often brightly spotted, as if to attract attention; but they can both emit fluids of a very disagreeable nature, they are certainly rejected by some birds, and are probably never eaten by any.

      The great family of ground beetles (Carabidæ) almost all possess a disagreeable and some a very pungent smell, and a few, called bombardier beetles, have the peculiar faculty of emitting a jet of very volatile liquid, which appears like a puff of smoke, and is accompanied by a distinct crepitating explosion. It is probably because these insects are mostly nocturnal and predacious that they do not present more vivid hues. They are chiefly remarkable for brilliant metallic tints or dull red patches when they are not wholly black, and are therefore very conspicuous by day, when insect-eaters are kept off by their bad odour and taste, but are sufficiently invisible at night when it is of importance that their prey should not become aware of their proximity.

      It seems probable that in some cases that which would appear at first to be a source of danger to its possessor may really be a means of protection. Many showy and weak-flying butterflies have a very broad expanse of wing, as in the brilliant blue Morphos of Brazilian forests, and the large Eastern Papilios; yet these groups are tolerably plentiful. Now, specimens of these butterflies are often captured with pierced and broken wings, as if they had been seized by birds from whom they had escaped; but if the wings had been much smaller in proportion to the body, it seems probable that the insect would be more frequently struck or pierced in a vital part, and thus the increased expanse of the wings may have been indirectly beneficial.

      In other cases the capacity of increase in a species is so great that however many of the perfect insect may be destroyed, there is always ample means for the continuance of the race. Many of the flesh flies, gnats, ants, palm-tree weevils and locusts are in this category. The whole family of Cetoniadæ or rose chafers, so full of gaily-coloured species, are probably saved from attack by a combination of characters. They fly very rapidly with a zigzag or waving course; they hide themselves the moment they alight, either in the corolla of flowers, or in rotten wood, or in cracks and hollows of trees, and they are generally encased in a very hard and polished coat of mail which may render them unsatisfactory food to such birds as would be able to capture them. The causes which lead to the development of colour have been here able to act unchecked, and we see the result in a large variety of the most gorgeously-coloured insects.

      Here, then, with our very imperfect knowledge of the life-history of animals, we are able to see that there are widely varied modes by which they may obtain protection from their enemies or concealment from their prey. Some of those seem to be so complete and effectual as to answer all the wants of the race, and lead to the maintenance of the largest possible population. When this is the case, we can well understand that no further protection derived from a modification of colour can be of the slightest use, and the most brilliant hues may be developed without any prejudicial effect upon the species. On some of the laws that determine the development of colour something may be said presently. It is now merely necessary to show that concealment by obscure or imitative tints is only one out of very many ways by which animals maintain their existence; and having done this we are prepared to consider the phenomena of what has been termed “mimicry.” It is to be particularly observed, however, that the word is not here used in the sense of voluntary imitation, but to imply a particular kind of resemblance—a resemblance not in internal structure but in external appearance—a resemblance in those parts only that catch the eye—a resemblance that deceives. As this kind of resemblance has the same effect as voluntary imitation or mimicry, and as we have no word that expresses the required meaning, “mimicry” was adopted by Mr. Bates (who was the first to explain the facts), and has led to some misunderstanding; but there need be none, if it is remembered that both “mimicry” and “imitation” are used in a metaphorical sense, as implying that close external likeness which causes things unlike in structure to be mistaken for each other.

      Mimicry

      It has been long known to entomologists that certain insects bear a strange external resemblance to others belonging to distinct genera, families, or even orders, and with which they have no real affinity whatever. The fact, however, appears to have been generally considered as dependent upon some unknown law of “analogy”—some “system of nature,” or “general plan,” which had guided the Creator in designing the myriads of insect forms, and which we could never hope to understand. In only one case does it appear that the resemblance was thought to be useful, and to have been designed as a means to a definite and intelligible purpose. The flies of the genus Volucella enter the nests of bees to deposit their eggs, so that their larvæ may feed upon the larvæ of the bees, and these flies are each wonderfully like the bee on which it is parasitic. Kirby and Spence believed that this resemblance or “mimicry” was for the express purpose of protecting the flies from the attacks of the bees, and the connection is so evident that it was hardly possible to avoid this conclusion. The resemblance, however, of moths to butterflies or to bees, of beetles to wasps, and of locusts to beetles, has been many times noticed by eminent writers; but scarcely ever till within the last few years does it appear to have been considered that these resemblances had any special purpose, or were of any direct benefit to the insects themselves. In this respect they were looked upon as accidental, as instances of the “curious analogies” in nature which must be wondered at but which could not be explained. Recently, however, these instances have been greatly multiplied; the nature of the resemblances has been more carefully studied, and it has been found that they are often carried out into such details as almost to imply a purpose of deceiving the observer. The phenomena, moreover, have been shown to follow certain definite laws, which again all indicate their dependence on the more general law of the “survival of the fittest,” or “the preservation of favoured races in the struggle for life.” It will, perhaps, be as well here to state what these laws or general conclusions are, and then to give some account of the facts which support them.

      The first law is, that in an overwhelming majority of cases of mimicry, the animals (or the groups) which resemble each other inhabit the same country, the same district, and in most cases are to be found together on the

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