In my ‘Origin of Species’[198] I briefly suggested that the long tail of the peacock would be inconvenient, and the conspicuous black colour of the male capercailzie dangerous, to the female during the period of incubation; and consequently that the transmission of these characters from the male to the female offspring had been checked through natural selection. I still think that this may have occurred in some few instances: but after mature reflection on all the facts which I have been able to collect, I am now inclined to believe that when the sexes differ, the successive variations have generally been from the first limited in their transmission to the same sex in which they first appeared. Since my remarks appeared, the subject of sexual coloration has been discussed in some very interesting papers by Mr. Wallace,[199] who believes that in almost all cases the successive variations tended at first to be transmitted equally to both sexes; but that the female was saved, through natural selection, from acquiring the conspicuous colours of the male, owing to the danger which she would thus have incurred during incubation.
This view necessitates a tedious discussion on a difficult point, namely whether the transmission of a character, which is at first inherited by both sexes, can be subsequently limited in its transmission, by means of selection, to one sex alone. We must bear in mind, as shewn in the preliminary chapter on sexual selection, that characters which are limited in their development to one sex are always latent in the other. An imaginary illustration will best aid us in seeing the difficulty of the case: we may suppose that a fancier wished to make a breed of pigeons, in which the males alone should be coloured of a pale blue, whilst the females retained their former slaty tint. As with pigeons characters of all kinds are usually transmitted to both sexes equally, the fancier would have to try to convert this latter form of inheritance into sexually-limited transmission. All that he could do would be to persevere in selecting every male pigeon which was in the least degree of a paler blue; and the natural result of this process, if steadily carried on for a long time, and if the pale variations were strongly inherited or often recurred, would be to make his whole stock of a lighter blue. But our fancier would be compelled to match, generation after generation, his pale blue males with slaty females, for he wishes to keep the latter of this colour. The result would generally be the production either of a mongrel piebald lot, or more probably the speedy and complete loss of the pale-blue colour, for the primordial slaty tint would be transmitted with prepotent force. Supposing, however, that some pale-blue males and slaty females were produced during each successive generation, and were always crossed together; then the slaty females would have, if I may use the expression, much blue blood in their veins, for their fathers, grandfathers, etc., will all have been blue birds. Under these circumstances it is conceivable (though I know of no distinct facts rendering it probable) that the slaty females might acquire so strong a latent tendency to pale-blueness, that they would not destroy this colour in their male offspring, their female offspring still inheriting the slaty tint. If so, the desired end of making a breed with the two sexes permanently different in colour might be gained.
The extreme importance, or rather necessity, of the desired character in the above case, namely, pale-blueness, being present though in a latent state in the female, so that the male offspring should not be deteriorated, will be best appreciated as follows: the male of Sœmmerring’s pheasant has a tail thirty-seven inches in length, whilst that of the female is only eight inches; the tail of the male common pheasant is about twenty inches, and that of the female twelve inches long. Now if the female Sœmmerring pheasant with her short tail were crossed with the male common pheasant, there can be no doubt that the male hybrid offspring would have a much longer tail than that of the pure offspring of the common pheasant. On the other hand, if the female common pheasant, with her tail nearly twice as long as that of the female Sœmmerring pheasant, were crossed with the male of the latter, the male hybrid offspring would have a much shorter tail than that of the pure offspring of Sœmmerring’s pheasant.[200]
Our fancier, in order to make his new breed with the males of a decided pale-blue tint, and the females unchanged, would have to continue selecting the males during many generations; and each stage of paleness would have to be fixed in the males, and rendered latent in the females. The task would be an extremely difficult one, and has never been tried, but might possibly succeed. The chief obstacle would be the early and complete loss of the pale-blue tint, from the necessity of reiterated crosses with the slaty female, the latter not having at first any latent tendency to produce pale-blue offspring.
On the other hand, if one or two males were to vary ever so slightly in paleness, and the variations were from the first limited in their transmission to the male sex, the task of making a new breed of the desired kind would be easy, for such males would simply have to be selected and matched with ordinary females. An analogous case has actually occurred, for there are breeds of the pigeon in Belgium[201] in which the males alone are marked with black striæ. In the case of the fowl, variations of colour limited in their transmission to the male sex habitually occur. Even when this form of inheritance prevails, it might well happen that some of the successive steps in the process of variation might be transferred to the female, who would then come to resemble in a slight degree the male, as occurs in some breeds of the fowl. Or again, the greater number, but not all, of the successive steps might be transferred to both sexes, and the female would then closely resemble the male. There can hardly be a doubt that this is the cause of the male pouter pigeon having a somewhat larger crop, and of the male carrier pigeon having somewhat larger wattles, than their respective females; for fanciers have not selected one sex more than the other, and have had no wish that these characters should be more strongly displayed in the male than in the female, yet this is the case with both breeds.
The same process would have to be followed, and the same difficulties would be encountered, if it were desired to make a breed with the females alone of some new colour.
Lastly, our fancier might wish to make a breed with the two sexes differing from each other, and both from the parent-species. Here the difficulty would be extreme, unless the successive variations were from the first sexually limited on both sides, and then there would be no difficulty. We see this with the fowl; thus the two sexes of the pencilled Hamburghs differ greatly from each other, and from the two sexes of the aboriginal Gallus bankiva; and both are now kept constant to their standard of excellence by continued selection, which would be impossible unless the distinctive characters of both were limited in their transmission. The Spanish fowl offers a more curious case; the male has an immense comb, but some of the successive variations, by the accumulation of which it was acquired, appear to have been transferred to the female; for she has a comb many times larger than that of the females of the parent-species. But the comb of the female differs in one respect from that of the male, for it is apt to lop over; and within a recent period it has been ordered by the fancy that this should always be the case, and success has quickly followed the order. Now the lopping of the comb must be sexually limited in its transmission, otherwise it would prevent the comb of the male from being perfectly upright, which would be abhorrent to every fancier. On the other hand the uprightness of the comb in the male must likewise be a sexually-limited character, otherwise it would prevent the comb of the female from lopping over.
From the foregoing illustrations, we see that even with almost unlimited time at command, it would be an extremely difficult and complex process, though perhaps not impossible, to change through selection one form of transmission into the other. Therefore, without distinct evidence in each case, I am unwilling to admit that this has often been effected with natural species. On the other hand by means of successive variations, which were from the first sexually limited in their transmission, there would not be the least difficulty in rendering a male bird widely different in colour or in any other character from the female; the latter being left unaltered, or slightly altered, or specially modified for the sake of protection.
As bright colours are of service to the males in their rivalry with other males, such colours would be selected, whether or not they were transmitted exclusively to the same sex. Consequently the females might be expected often to partake of the brightness of the males to a greater or less degree; and this occurs with a host of species. If all the successive variations were transmitted equally to both sexes, the females would be undistinguishable from the males; and this likewise occurs with many birds. If, however, dull colours were of high importance for the safety of the female during incubation, as with many ground birds, the females which varied in brightness, or which received through inheritance from the males any marked accession of brightness, would sooner or later be destroyed. But the tendency in the males to continue for an indefinite period transmitting to their female offspring their own brightness, would have to be eliminated by a change in the form of inheritance; and this, as shewn by our previous illustration, would be extremely difficult. The more probable result of the long-continued destruction of the more brightly-coloured females, supposing the equal form of transmission to prevail, would be the lessening or annihilation of the bright colours of the males, owing to their continually crossing with the duller females. It would be tedious to follow out all the other possible results; but I may remind the reader, as shewn in the eighth chapter, that if sexually-limited variations in brightness occurred in the females, even if they were not in the least injurious to them and consequently were not eliminated, yet they would not be favoured or selected, for the male usually accepts any female, and does not select the more attractive individuals; consequently these variations would be liable to be lost, and would have little influence on the character of the race; and this will aid in accounting for the females being commonly less brightly-coloured than the males.
In the chapter just referred to, instances were given, and any number might have been added, of variations occurring at different ages, and inherited at the same age. It was also shewn that variations which occur late in life are commonly transmitted to the same sex in which they first appeared; whilst variations occurring early in life are apt to be transmitted to both sexes; not that all the cases of sexually-limited transmission can thus be accounted for. It was further shewn that if a male bird varied by becoming brighter whilst young, such variations would be of no service until the age for reproduction had arrived, and there was competition between rival males. If we suppose that three-fourths of the young males of any species are on an average destroyed by various enemies; then the chances would be as three to one against any one individual more brightly-coloured than usual surviving to propagate its kind. But in the case of birds which live on the ground and which commonly need the protection of dull colours, bright tints would be far more dangerous to the young and inexperienced than to the adult males. Consequently the males which varied in brightness whilst young would suffer much destruction and be eliminated through natural selection; on the other hand the males which varied in this manner when nearly mature, notwithstanding that they were exposed to some additional danger, might survive, and from being favoured through sexual selection, would procreate their kind. The brightly-coloured young males being destroyed and the mature ones being successful in their courtship, may account, on the principle of a relation existing between the period of variation and the form of transmission, for the males alone of many birds, having acquired and transmitted brilliant colours to their male offspring alone. But I by no means wish to maintain that the influence of age on the form of transmission is indirectly the sole cause of the great difference in brilliancy between the sexes of many birds.