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Mr. Wallace remarks, "distastefulness alone would be insufficient "to protect a caterpillar unless some outward sign indicated to "its would-be destroyer that its prey was a disgusting morsel." Under these circumstances it would be highly advantageous to a caterpillar to be instantaneously and certainly recognised as unpalatable by all birds and other animals. Thus the most gaudy colours would be serviceable, and might have been · gained by variation and the survival of the most easily-recognised individuals.

33

This hypothesis appears at first sight very bold, but when it was brought before the Entomological Society & it was supported by various statements; and Mr. J. Jenner Weir, who keeps a large number of birds in an aviary, informs me that he has made many trials, and finds no exception to the rule, that ali caterpillars of nocturnal and retiring habits with smooth skins, all of a green colour, and all which imitate twigs, are greedily devoured by his birds. The hairy and spinose kinds are invariably rejected, as were four conspicuously-coloured species. When the birds rejected a caterpillar, they plainly shewed, by shaking their heads, and cleansing their beaks, that they were disgusted by the taste. Three conspicuous kinds of caterpillars and moths were also given to some lizards and frogs, by Mr. A. Butler, and were rejected, though other kinds were eagerly eaten. Thus the probability of Mr. Wallace's view is confirmed, namely, that certain caterpillars have been made conspicuous for their own good, so as to be easily recognised by their enemies, on nearly the same principle that poisons are sold in coloured bottles by druggists for the good of man. We cannot, however, at present thus explain the elegant diversity in the colours of many caterpillars; but any species which had at some former period acquired a dull, mottled, or striped appearance, either in imitation of surrounding objects, or from the direct action of climate, &c., almost certainly would not become uniform in colour, when its tints were rendered intense and bright; for in order to make a caterpillar merely conspicuous, there would be no selection in any definite direction.

Summary and Concluding Remarks on Insects.-Looking back 83 Proc. Entomolog. Soc.' Dec. 3rd, 1866, p. xlv., and March 4th, 1867, p. lxxx.

34 See Mr. J. Jenner Weir's paper on Insects and Insectivorous Birds, in Transact. Ent. Soc.' 1869, p. 21; also Mr. Butler's paper, ibid. p. 27. Mr. Riley has given

analogous facts in the Third Annual Report on the Noxious Insects of Missouri,' 1871, p. 148. Some opposed cases are, however, given by Dr. Wallace and M. H. d'Orville; see Zoological Record,' 1869, F.

349,

to the several Orders, we see that the sexes often differ in various characters, the meaning of which is not in the least understood. The sexes, also, often differ in their organs of sense and means of locomotion, so that the males may quickly discover and reach the females. They differ still oftener in the males possessing diversified contrivances for retaining the females when found. We are, however, here concerned only in a secondary degree with sexual differences of these kinds.

In almost all the Orders, the males of some species, even of weak and delicate kinds, are known to be highly pugnacious; and some few are furnished with special weapons for fighting with their rivals. But the law of battle does not prevail nearly so widely with insects as with the higher animals. Hence it probably arises, that it is in only a few cases that the males have been rendered larger and stronger than the females. On the contrary, they are usually smaller, so that they may be developed within a shorter time, to be ready in large numbers for the emergence of the females.

In two families of the Homoptera and in three of the Orthoptera, the males alone possess sound-producing organs in an efficient state. These are used incessantly during the breedingseason, not only for calling the females, but apparently for charming or exciting them in rivalry with other males. No one who admits the agency of selection of any kind, will, after reading the above discussion, dispute that these musical instruments have been acquired through sexual selection. In four other Orders the members of one sex, or more commonly of both sexes, are provided with organs for producing various sounds, which apparently serve merely as call-notes. When both sexes are thus provided, the individuals which were able to make the loudest or most continuous noise would gain partners before those which were less noisy, so that their organs have probably been gained through sexual selection. It is instructive to reflect on the wonderful diversity of the means for producing sound, possessed by the males alone, or by both sexes, in no less than six Orders. We thus learn how effectual sexual selection has been in leading to modifications which sometimes, as with the Homoptera, relate to important parts of the organisation.

From the reasons assigned in the last chapter, it is probable that the great horns possessed by the males of many Lamellicorn, and some other beetles, have been acquired as ornaments. From the small size of insects, we are apt to undervalue their appearance. If we could imagine a male Chalcosoma (fig. 16), with its polished bronzed coat of mail, and its vast complex

horns, magnified to the size of a horse, or even of a dog, it would be one of the most imposing animals in the world.

The colouring of insects is a complex and obscure subject. When the male differs slightly from the female, and neither are brilliantly-coloured, it is probable that the sexes have varied in a slightly different manner, and that the variations have been transmitted by each sex to the same, without any benefit or evil thus accruing. When the male is brilliantly-coloured and differs conspicuously from the female, as with some dragon-flies and many butterflies, it is probable that he owes his colours to sexual selection; whilst the female has retained a primordial or very ancient type of colouring, slightly modified by the agencies before explained. But in some cases the female has apparently been made obscure by variations transmitted to her alone, as a means of direct protection; and it is almost certain that she has sometimes been made brilliant, so as to imitate other protected species inhabiting the same district. When the sexes resemble each other and both are obscurely coloured, there is no doubt that they have been in a multitude of cases so coloured for the sake of protection. So it is in some instances when both are brightly-coloured, for they thus imitate protected species, or resemble surrounding objects such as flowers; or they give potice to their enemies that they are unpalatable. In other cases in which the sexes resemble each other and are both brilliant, especially when the colours are arranged for display, we may conclude that they have been gained by the male sex as an attraction, and have been transferred to the female. We are more especially led to this conclusion whenever the same type of coloration prevails throughout a whole group, and we find that the males of some species differ widely in colour from the females, whilst others differ slightly or not at all, with intermediate gradations connecting these extreme states.

In the same manner as bright colours have often been partially transferred from the males to the females, so it has been with the extraordinary horns of many Lamellicorn and some other beetles. So again, the sound-producing organs proper to the males of the Homoptera and Orthoptera have generally been transferred in a rudimentary, or even in a nearly perfect condition, to the females; yet not sufficiently perfect to be of any use. It is also an interesting fact, as bearing on sexual selection, that the stridulating organs of certain male Orthoptera are not fully developed until the last moult; and that the colours of certain male dragon-flies are not fully developed until some little time after their emergence from the pupal state, and when they are ready to breed.

Sexual selection implies that the more attractive individuals are preferred by the opposite sex; and as with insects, when tne sexes differ, it is the male which, with some rare exceptions, is the more ornamented, and departs more from the type to which the species belongs;--and as it is the male which searches eagerly for the female, we must suppose that the females habitually or occasionally prefer the more beautiful males, and that these have thus acquired their beauty. That the females in most or all the orders would have the power of rejecting any particular male, is probable from the many singular contrivances possessed by the males, such as great jaws, adhesive cushions, spines, elongated legs, &c., for seizing the female; for these contrivances shew that there is some difficulty in the act, so that her concurrence would seem necessary. Judging from what we know of the perceptive powers and affections of various insects, there is no antecedent improbability in sexual selection having come largely into play; but we have as yet no direct evidence on this head, and some facts are opposed to the belief. Nevertheless, when we see many males pursuing the same female, we can hardly believe that the pairing is left to blind chance that the female exerts no choice, and is not influenced by the gorgeous colours or other ornaments with which the male is decorated.

If we admit that the females of the Homoptera and Orthoptera appreciate the musical tones of their male partners, and that the various instruments have been perfected through sexual selection, there is little improbability in the females of other insects appreciating beauty in form or colour, and consequently in such characters having been thus gained by the males. But from the circumstance of colour being so variable, and from its having been so often modified for the sake of protection, it is difficult to decide in how large a proportion of cases sexual selection has played a part. This is more especially difficult in those Orders, such as Orthoptera, Hymenoptera, and Coleoptera, in which the two sexes rarely differ much in colour; for we are then left to mere analogy. With the Coleoptera, however, as before remarked, it is in the great Lamellicorn group, placed by some authors at the head of the Order, and in which we sometimes see a mutual attachment between the sexes, that we find the males of some species possessing weapons for sexual strife, others furnished with wonderful horns, many with stridulating organs, and others ornamented with splendid metallic tints. Hence it seems probable that all these characters have been gained through the same means, namely sexual selection. With butterflies we have the best evidence, as the males

sometimes take pains to display their beautiful colours; and we cannot believe that they would act thus, unless the display was of use to them in their courtship.

When we treat of Birds, we shall see that they present in their secondary sexual characters the closest analogy with insects. Thus, many male birds are highly pugnacious, and some are furnished with special weapons for fighting with their rivals. They possess organs which are used during the breedingseason for producing vocal and instrumental music. They are frequently ornamented with combs, horns, wattles and plumes of the most diversified kinds, and are decorated with beautiful colours, all evidently for the sake of display. We shall find that, as with insects, both sexes in certain groups are equally beautiful, and are equally provided with ornaments which are usually confined to the male sex. In other groups both sexes are equally plain-coloured and unornamented. Lastly, in some few anomalous cases, the females are more beautiful than the males. We shall often find, in the same group of birds, every gradation from no difference between the sexes, to an extreme difference. We shall see that female birds, like female insects, often possess more or less plain traces or rudiments of characters which properly belong to the males and are of use only to them. The analogy, indeed, in all these respects between birds and insects is curiously close. Whatever explanation applies to the one class probably applies to the other; and this explanation, as we shall hereafter attempt to shew in further detail, is sexual selection.

CHAPTER XII.

SECONDARY SEXUAL CHARACTERS OF FISHES, AMPHIBIANS,

AND REPTILES.

FISHES: Courtship and battles of the males-Larger size of the females -Males, bright colours and ornamental appendages; other strange characters-Colours and appendages acquired by the males durind the breeding-season alone-Fishes with both sexes brilliantly coloured -Protective colours-The less conspicuous colours of the female cannot be accounted for on the principle of protection-Male fishes building nests, and taking charge of the ova and young. AMPHIBIANS: Differences in structure and colour between the sexes-Vocal organs. REPTILES: Chelonians-Crocodiles-Snakes, colours in some cases protective—Lizards, battles of-Ornamental appendages-Strange differences in structure between the sexes-Colours-Sexual differences almost as great as with birds.

We have now arrived at the great sub-kingdom of the Vertebrata, and will commence with the lowest class, that of Fishes. The.

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