The Descent of Man, and Selection in Relation to Sex

I have as yet only referred to the species in which the males are brighter coloured than the females, and I have attributed their beauty to the females for many generations having chosen and paired with the more attractive males. But converse cases occur, though rarely, in which the females are more brilliant than the males; and here, as I believe, the males have selected the more beautiful females, and have thus slowly added to their beauty. We do not know why in various classes of animals the males of some few species have selected the more beautiful females instead of having gladly accepted any female, as seems to be the general rule in the animal kingdom: but if, contrary to what generally occurs with the Lepidoptera, the females were much more numerous than the males, the latter would be likely to pick out the more beautiful females. Mr. Butler shewed me several species of Callidryas in the British Museum, in some of which the females equalled, and in others greatly surpassed the males in beauty; for the females alone have the borders of their wings suffused with crimson and orange, and spotted with black. The plainer males of these species closely resemble each other, shewing that here the females have been modified; whereas in those cases, where the males are the more ornate, it is these which have been modified, the females remaining closely alike.

In England we have some analogous cases, though not so marked. The females alone of two species of Thecla have a bright-purple or orange patch on their fore-wings. In Hipparchia the sexes do not differ much; but it is the female of H. janira which has a conspicuous light-brown patch on her wings; and the females of some of the other species are brighter coloured than their males. Again, the females of Colias edusa and hyale have "orange or yellow spots on the black marginal border, represented in the males only by thin streaks"; and in Pieris it is the females which "are ornamented with black spots on the fore-wings, and these are only partially present in the males." Now the males of many butterflies are known to support the females during their marriage flight; but in the species just named it is the females which support the males; so that the part which the two sexes play is reversed, as is their relative beauty. Throughout the animal kingdom the males commonly take the more active share in wooing, and their beauty seems to have been increased by the females having accepted the more attractive individuals; but with these butterflies, the females take the more active part in the final marriage ceremony, so that we may suppose that they likewise do so in the wooing; and in this case we can understand how it is that they have been rendered the more beautiful. Mr. Meldola, from whom the foregoing statements have been taken, says in conclusion: "Though I am not convinced of the action of sexual selection in producing the colours of insects, it cannot be denied that these facts are strikingly corroborative of Mr. Darwin's views." (23. 'Nature,' April 27, 1871, p. 508. Mr. Meldola quotes Donzel, in 'Soc. Ent. de France,' 1837, p. 77, on the flight of butterflies whilst pairing. See also Mr. G. Fraser, in 'Nature,' April 20, 1871, p. 489, on the sexual differences of several British butterflies.)

As sexual selection primarily depends on variability, a few words must be added on this subject. In respect to colour there is no difficulty, for any number of highly variable Lepidoptera could be named. One good instance will suffice. Mr. Bates shewed me a whole series of specimens of Papilio sesostris and P. childrenae; in the latter the males varied much in the extent of the beautifully enamelled green patch on the fore-wings, and in the size of the white mark, and of the splendid crimson stripe on the hind-wings; so that there was a great contrast amongst the males between the most and the least gaudy. The male of Papilio sesostris is much less beautiful than of P. childrenae; and it likewise varies a little in the size of the green patch on the fore-wings, and in the occasional appearance of the small crimson stripe on the hind-wings, borrowed, as it would seem, from its own female; for the females of this and of many other species in the Aeneas group possess this crimson stripe. Hence between the brightest specimens of P. sesostris and the dullest of P. childrenae, there was but a small interval; and it was evident that as far as mere variability is concerned, there would be no difficulty in permanently increasing the beauty of either species by means of selection. The variability is here almost confined to the male sex; but Mr. Wallace and Mr. Bates have shewn (24. Wallace on the Papilionidae of the Malayan Region, in 'Transact. Linn. Soc.' vol. xxv. 1865, pp. 8, 36. A striking case of a rare variety, strictly intermediate between two other well-marked female varieties, is given by Mr. Wallace. See also Mr. Bates, in 'Proc. Entomolog. Soc.' Nov. 19, 1866, p. xl.) that the females of some species are extremely variable, the males being nearly constant. In a future chapter I shall have occasion to shew that the beautiful eye-like spots, or ocelli, found on the wings of many Lepidoptera, are eminently variable. I may here add that these ocelli offer a difficulty on the theory of sexual selection; for though appearing to us so ornamental, they are never present in one sex and absent in the other, nor do they ever differ much in the two sexes. (25. Mr. Bates was so kind as to lay this subject before the Entomological Society, and I have received answers to this effect from several entomologists.) This fact is at present inexplicable; but if it should hereafter be found that the formation of an ocellus is due to some change in the tissues of the wings, for instance, occurring at a very early period of development, we might expect, from what we know of the laws of inheritance, that it would be transmitted to both sexes, though arising and perfected in one sex alone.

On the whole, although many serious objections may be urged, it seems probable that most of the brilliantly-coloured species of Lepidoptera owe their colours to sexual selection, excepting in certain cases, presently to be mentioned, in which conspicuous colours have been gained through mimicry as a protection. From the ardour of the male throughout the animal kingdom, he is generally willing to accept any female; and it is the female which usually exerts a choice. Hence, if sexual selection has been efficient with the Lepidoptera, the male, when the sexes differ, ought to be the more brilliantly coloured, and this undoubtedly is the case. When both sexes are brilliantly coloured and resemble each other, the characters acquired by the males appear to have been transmitted to both. We are led to this conclusion by cases, even within the same genus, of gradation from an extraordinary amount of difference to identity in colour between the two sexes.

But it may be asked whether the difference in colour between the sexes may not be accounted for by other means besides sexual selection. Thus the males and females of the same species of butterfly are in several cases known (26. H.W. Bates, 'The Naturalist on the Amazons,' vol. ii. 1863, p. 228. A.R. Wallace, in 'Transactions, Linnean Society,' vol. xxv. 1865, p. 10.) to inhabit different stations, the former commonly basking in the sunshine, the latter haunting gloomy forests. It is therefore possible that different conditions of life may have acted directly on the two sexes; but this is not probable (27. On this whole subject see 'The Variation of Animals and Plants under Domestication,' 1868, vol. ii. chap. xxiii.) as in the adult state they are exposed to different conditions during a very short period; and the larvae of both are exposed to the same conditions. Mr. Wallace believes that the difference between the sexes is due not so much to the males having been modified, as to the females having in all or almost all cases acquired dull colours for the sake of protection. It seems to me, on the contrary, far more probable that it is the males which have been chiefly modified through sexual selection, the females having been comparatively little changed. We can thus understand how it is that the females of allied species generally resemble one another so much more closely than do the males. They thus shew us approximately the primordial colouring of the parent-species of the group to which they belong. They have, however, almost always been somewhat modified by the transfer to them of some of the successive variations, through the accumulation of which the males were rendered beautiful. But I do not wish to deny that the females alone of some species may have been specially modified for protection. In most cases the males and females of distinct species will have been exposed during their prolonged larval state to different conditions, and may have been thus affected; though with the males any slight change of colour thus caused will generally have been masked by the brilliant tints gained through sexual selection. When we treat of Birds, I shall have to discuss the whole question, as to how far the differences in colour between the sexes are due to the males having been modified through sexual selection for ornamental purposes, or to the females having been modified through natural selection for the sake of protection, so that I will here say but little on the subject.

In all the cases in which the more common form of equal inheritance by both sexes has prevailed, the selection of bright-coloured males would tend to make the females bright-coloured; and the selection of dull-coloured females would tend to make the males dull. If both processes were carried on simultaneously, they would tend to counteract each other; and the final result would depend on whether a greater number of females from being well protected by obscure colours, or a greater number of males by being brightly-coloured and thus finding partners, succeeded in leaving more numerous offspring.

In order to account for the frequent transmission of characters to one sex alone, Mr. Wallace expresses his belief that the more common form of equal inheritance by both sexes can be changed through natural selection into inheritance by one sex alone, but in favour of this view I can discover no evidence. We know from what occurs under domestication that new characters often appear, which from the first are transmitted to one sex alone; and by the selection of such variations there would not be the slightest difficulty in giving bright colours to the males alone, and at the same time or subsequently, dull colours to the females alone. In this manner the females of some butterflies and moths have, it is probable, been rendered inconspicuous for the sake of protection, and widely different from their males.

I am, however, unwilling without distinct evidence to admit that two complex processes of selection, each requiring the transference of new characters to one sex alone, have been carried on with a multitude of species, – that the males have been rendered more brilliant by beating their rivals, and the females more dull-coloured by having escaped from their enemies. The male, for instance, of the common brimstone butterfly (Gonepteryx), is of a far more intense yellow than the female, though she is equally conspicuous; and it does not seem probable that she specially acquired her pale tints as a protection, though it is probable that the male acquired his bright colours as a sexual attraction. The female of Anthocharis cardamines does not possess the beautiful orange wing-tips of the male; consequently she closely resembles the white butterflies (Pieris) so common in our gardens; but we have no evidence that this resemblance is beneficial to her. As, on the other hand, she resembles both sexes of several other species of the genus inhabiting various quarters of the world, it is probable that she has simply retained to a large extent her primordial colours.

Finally, as we have seen, various considerations lead to the conclusion that with the greater number of brilliantly-coloured Lepidoptera it is the male which has been chiefly modified through sexual selection; the amount of difference between the sexes mostly depending on the form of inheritance which has prevailed. Inheritance is governed by so many unknown laws or conditions, that it seems to us to act in a capricious manner (28. The 'Variation of Animals and Plants under Domestication,' vol. ii. chap. xii. p. 17.); and we can thus, to a certain extent, understand how it is that with closely allied species the sexes either differ to an astonishing degree, or are identical in colour. As all the successive steps in the process of variation are necessarily transmitted through the female, a greater or less number of such steps might readily become developed in her; and thus we can understand the frequent gradations from an extreme difference to none at all between the sexes of allied species. These cases of gradation, it may be added, are much too common to favour the supposition that we here see females actually undergoing the process of transition and losing their brightness for the sake of protection; for we have every reason to conclude that at any one time the greater number of species are in a fixed condition.

MIMICRY

This principle was first made clear in an admirable paper by Mr. Bates (29. 'Transact. Linn. Soc.' vol. xxiii. 1862, p. 495.), who thus threw a flood of light on many obscure problems. It had previously been observed that certain butterflies in S. America belonging to quite distinct families, resembled the Heliconidae so closely in every stripe and shade of colour, that they could not be distinguished save by an experienced entomologist. As the Heliconidae are coloured in their usual manner, whilst the others depart from the usual colouring of the groups to which they belong, it is clear that the latter are the imitators, and the Heliconidae the imitated. Mr. Bates further observed that the imitating species are comparatively rare, whilst the imitated abound, and that the two sets live mingled together. From the fact of the Heliconidae being conspicuous and beautiful insects, yet so numerous in individuals and species, he concluded that they must be protected from the attacks of enemies by some secretion or odour; and this conclusion has now been amply confirmed (30. 'Proc. Entomological Soc.' Dec. 3, 1866, p. xlv.), especially by Mr. Belt. Hence Mr. Bates inferred that the butterflies which imitate the protected species have acquired their present marvellously deceptive appearance through variation and natural selection, in order to be mistaken for the protected kinds, and thus to escape being devoured. No explanation is here attempted of the brilliant colours of the imitated, but only of the imitating butterflies. We must account for the colours of the former in the same general manner, as in the cases previously discussed in this chapter. Since the publication of Mr. Bates' paper, similar and equally striking facts have been observed by Mr. Wallace in the Malayan region, by Mr. Trimen in South Africa, and by Mr. Riley in the United States. (31. Wallace, 'Transact. Linn. Soc.' vol. xxv. 1865 p. i.; also, 'Transact. Ent. Soc.' vol. iv. (3rd series), 1867, p. 301. Trimen, 'Linn. Transact.' vol. xxvi. 1869, p. 497. Riley, 'Third Annual Report on the Noxious Insects of Missouri,' 1871, pp. 163-168. This latter essay is valuable, as Mr. Riley here discusses all the objections which have been raised against Mr. Bates's theory.)

As some writers have felt much difficulty in understanding how the first steps in the process of mimicry could have been effected through natural selection, it may be well to remark that the process probably commenced long ago between forms not widely dissimilar in colour. In this case even a slight variation would be beneficial, if it rendered the one species more like the other; and afterwards the imitated species might be modified to an extreme degree through sexual selection or other means, and if the changes were gradual, the imitators might easily be led along the same track, until they differed to an equally extreme degree from their original condition; and they would thus ultimately assume an appearance or colouring wholly unlike that of the other members of the group to which they belonged. It should also be remembered that many species of Lepidoptera are liable to considerable and abrupt variations in colour. A few instances have been given in this chapter; and many more may be found in the papers of Mr. Bates and Mr. Wallace.

With several species the sexes are alike, and imitate the two sexes of another species. But Mr. Trimen gives, in the paper already referred to, three cases in which the sexes of the imitated form differ from each other in colour, and the sexes of the imitating form differ in a like manner. Several cases have also been recorded where the females alone imitate brilliantly-coloured and protected species, the males retaining "the normal aspect of their immediate congeners." It is here obvious that the successive variations by which the female has been modified have been transmitted to her alone. It is, however, probable that some of the many successive variations would have been transmitted to, and developed in, the males had not such males been eliminated by being thus rendered less attractive to the females; so that only those variations were preserved which were from the first strictly limited in their transmission to the female sex. We have a partial illustration of these remarks in a statement by Mr. Belt (32. 'The Naturalist in Nicaragua,' 1874, p. 385.); that the males of some of the Leptalides, which imitate protected species, still retain in a concealed manner some of their original characters. Thus in the males "the upper half of the lower wing is of a pure white, whilst all the rest of the wings is barred and spotted with black, red and yellow, like the species they mimic. The females have not this white patch, and the males usually conceal it by covering it with the upper wing, so that I cannot imagine its being of any other use to them than as an attraction in courtship, when they exhibit it to the females, and thus gratify their deep-seated preference for the normal colour of the Order to which the Leptalides belong."

BRIGHT COLOURS OF CATERPILLARS

Whilst reflecting on the beauty of many butterflies, it occurred to me that some caterpillars were splendidly coloured; and as sexual selection could not possibly have here acted, it appeared rash to attribute the beauty of the mature insect to this agency, unless the bright colours of their larvae could be somehow explained. In the first place, it may be observed that the colours of caterpillars do not stand in any close correlation with those of the mature insect. Secondly, their bright colours do not serve in any ordinary manner as a protection. Mr. Bates informs me, as an instance of this, that the most conspicuous caterpillar which he ever beheld (that of a Sphinx) lived on the large green leaves of a tree on the open llanos of South America; it was about four inches in length, transversely banded with black and yellow, and with its head, legs, and tail of a bright red. Hence it caught the eye of any one who passed by, even at the distance of many yards, and no doubt that of every passing bird.

I then applied to Mr. Wallace, who has an innate genius for solving difficulties. After some consideration he replied: "Most caterpillars require protection, as may be inferred from some kinds being furnished with spines or irritating hairs, and from many being coloured green like the leaves on which they feed, or being curiously like the twigs of the trees on which they live." Another instance of protection, furnished me by Mr. J. Mansel Weale, may be added, namely, that there is a caterpillar of a moth which lives on the mimosas in South Africa, and fabricates for itself a case quite indistinguishable from the surrounding thorns. From such considerations Mr. Wallace thought it probable that conspicuously coloured caterpillars were protected by having a nauseous taste; but as their skin is extremely tender, and as their intestines readily protrude from a wound, a slight peck from the beak of a bird would be as fatal to them as if they had been devoured. Hence, as 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.

This hypothesis appears at first sight very bold, but when it was brought before the Entomological Society (33. 'Proceedings, Entomological Society,' Dec. 3, 1866, p. xlv. and March 4, 1867, p. lxxx.) 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 all 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. (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, p. 349.) 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, etc., 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 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 breeding-season, 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 notice 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 the 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, etc., for seizing the female; for these contrivances show 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.