We may now consider Mr. Wallace’s arguments in regard to the sexual coloration of birds. He believes that the bright tints originally acquired through sexual selection by the males would in all, or almost all cases, have been transmitted to the females, unless the transference had been checked through natural selection. I may here remind the reader that various facts opposed to this view have already been given under reptiles, amphibians, fishes and lepidoptera. Mr. Wallace rests his belief chiefly, but not exclusively, as we shall see in the next chapter, on the following statement (9. ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 78.), that when both sexes are coloured in a very conspicuous manner, the nest is of such a nature as to conceal the sitting bird; but when there is a marked contrast of colour between the sexes, the male being gay and the female dull-coloured, the nest is open and exposes the sitting bird to view. This coincidence, as far as it goes, certainly seems to favour the belief that the females which sit on open nests have been specially modified for the sake of protection; but we shall presently see that there is another and more probable explanation, namely, that conspicuous females have acquired the instinct of building domed nests oftener than dull-coloured birds. Mr. Wallace admits that there are, as might have been expected, some exceptions to his two rules, but it is a question whether the exceptions are not so numerous as seriously to invalidate them.
There is in the first place much truth in the Duke of Argyll’s remark (10. ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 281.) that a large domed nest is more conspicuous to an enemy, especially to all tree-haunting carnivorous animals, than a smaller open nest. Nor must we forget that with many birds which build open nests, the male sits on the eggs and aids the female in feeding the young: this is the case, for instance, with Pyranga aestiva (11. Audubon, ‘Ornithological Biography,’ vol. i. p. 233.), one of the most splendid birds in the United States, the male being vermilion, and the female light brownish-green. Now if brilliant colours had been extremely dangerous to birds whilst sitting on their open nests, the males in these cases would have suffered greatly. It might, however, be of such paramount importance to the male to be brilliantly coloured, in order to beat his rivals, that this may have more than compensated some additional danger.
Mr. Wallace admits that with the King-crows (Dicrurus), Orioles, and Pittidae, the females are conspicuously coloured, yet build open nests; but he urges that the birds of the first group are highly pugnacious and could defend themselves; that those of the second group take extreme care in concealing their open nests, but this does not invariably hold good (12. Jerdon, ‘Birds of India,’ vol. ii. p. 108. Gould’s ‘Handbook of the Birds of Australia,’ vol. i. p. 463.); and that with the birds of the third group the females are brightly coloured chiefly on the under surface. Besides these cases, pigeons which are sometimes brightly, and almost always conspicuously coloured, and which are notoriously liable to the attacks of birds of prey, offer a serious exception to the rule, for they almost always build open and exposed nests. In another large family, that of the humming-birds, all the species build open nests, yet with some of the most gorgeous species the sexes are alike; and in the majority, the females, though less brilliant than the males, are brightly coloured. Nor can it be maintained that all female humming-birds, which are brightly coloured, escape detection by their tints being green, for some display on their upper surfaces red, blue, and other colours. (13. For instance, the female Eupetomena macroura has the head and tail dark blue with reddish loins; the female Lampornis porphyrurus is blackish-green on the upper surface, with the lores and sides of the throat crimson; the female Eulampis jugularis has the top of the head and back green, but the loins and the tail are crimson. Many other instances of highly conspicuous females could be given. See Mr. Gould’s magnificent work on this family.)
In regard to birds which build in holes or construct domed nests, other advantages, as Mr. Wallace remarks, besides concealment are gained, such as shelter from the rain, greater warmth, and in hot countries protection from the sun (14. Mr. Salvin noticed in Guatemala (‘Ibis,’ 1864, p. 375) that humming-birds were much more unwilling to leave their nests during very hot weather, when the sun was shining brightly, as if their eggs would be thus injured, than during cool, cloudy, or rainy weather.); so that it is no valid objection to his view that many birds having both sexes obscurely coloured build concealed nests. (15. I may specify, as instances of dull-coloured birds building concealed nests, the species belonging to eight Australian genera described in Gould’s ‘Handbook of the Birds of Australia,’ vol. i. pp. 340, 362, 365, 383, 387, 389, 391, 414.) The female Horn-bill (Buceros), for instance, of India and Africa is protected during incubation with extraordinary care, for she plasters up with her own excrement the orifice of the hole in which she sits on her eggs, leaving only a small orifice through which the male feeds her; she is thus kept a close prisoner during the whole period of incubation (16. Mr. C. Horne, ‘Proc. Zoolog. Soc.’ 1869. p. 243.); yet female horn-bills are not more conspicuously coloured than many other birds of equal size which build open nests. It is a more serious objection to Mr. Wallace’s view, as is admitted by him, that in some few groups the males are brilliantly coloured and the females obscure, and yet the latter hatch their eggs in domed nests. This is the case with the Grallinae of Australia, the Superb Warblers (Maluridae) of the same country, the Sun-birds (Nectariniae), and with several of the Australian Honey-suckers or Meliphagidae. (17. On the nidification and colours of these latter species, see Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 504, 527.)
If we look to the birds of England we shall see that there is no close and general relation between the colours of the female and the nature of the nest which is constructed. About forty of our British birds (excluding those of large size which could defend themselves) build in holes in banks, rocks, or trees, or construct domed nests. If we take the colours of the female goldfinch, bullfinch, or blackbird, as a standard of the degree of conspicuousness, which is not highly dangerous to the sitting female, then out of the above forty birds the females of only twelve can be considered as conspicuous to a dangerous degree, the remaining twenty-eight being inconspicuous. (18. I have consulted, on this subject, Macgillivray’s ‘British Birds,’ and though doubts may be entertained in some cases in regard to the degree of concealment of the nest, and to the degree of conspicuousness of the female, yet the following birds, which all lay their eggs in holes or in domed nests, can hardly be considered, by the above standard, as conspicuous: Passer, 2 species; Sturnus, of which the female is considerably less brilliant than the male; Cinclus; Motallica boarula (?); Erithacus (?); Fruticola, 2 sp.; Saxicola; Ruticilla, 2 sp.; Sylvia, 3 sp.; Parus, 3 sp.; Mecistura; Anorthura; Certhia; Sitta; Yunx; Muscicapa, 2 sp.; Hirundo, 3 sp.; and Cypselus. The females of the following 12 birds may be considered as conspicuous according to the same standard, viz., Pastor, Motacilla alba, Parus major and P. caeruleus, Upupa, Picus, 4 sp., Coracias, Alcedo, and Merops.) Nor is there any close relation within the same genus between a well-pronounced difference in colour between the sexes, and the nature of the nest constructed. Thus the male house sparrow (Passer domesticus) differs much from the female, the male tree-sparrow (P. montanus) hardly at all, and yet both build well-concealed nests. The two sexes of the common fly-catcher (Muscicapa grisola) can hardly be distinguished, whilst the sexes of the pied fly-catcher (M. luctuosa) differ considerably, and both species build in holes or conceal their nests. The female blackbird (Turdus merula) differs much, the female ring-ouzel (T. torquatus) differs less, and the female common thrush (T. musicus) hardly at all from their respective males; yet all build open nests. On the other hand, the not very distantly-allied water-ouzel (Cinclus aquaticus) builds a domed nest, and the sexes differ about as much as in the ring-ouzel. The black and red grouse (Tetrao tetrix and T. scoticus) build open nests in equally well-concealed spots, but in the one species the sexes differ greatly, and in the other very little.
Notwithstanding the foregoing objections, I cannot doubt, after reading Mr. Wallace’s excellent essay, that looking to the birds of the world, a large majority of the species in which the females are conspicuously coloured (and in this case the males with rare exceptions are equally conspicuous), build concealed nests for the sake of protection. Mr. Wallace enumerates (19. ‘Journal of Travel,’ edited by A. Murray, vol. i. p. 78.) a long series of groups in which this rule holds good; but it will suffice here to give, as instances, the more familiar groups of kingfishers, toucans, trogons, puff-birds (Capitonidae), plantain-eaters (Musophagae, woodpeckers, and parrots. Mr. Wallace believes that in these groups, as the males gradually acquired through sexual selection their brilliant colours, these were transferred to the females and were not eliminated by natural selection, owing to the protection which they already enjoyed from their manner of nidification. According to this view, their present manner of nesting was acquired before their present colours. But it seems to me much more probable that in most cases, as the females were gradually rendered more and more brilliant from partaking of the colours of the male, they were gradually led to change their instincts (supposing that they originally built open nests), and to seek protection by building domed or concealed nests. No one who studies, for instance, Audubon’s account of the differences in the nests of the same species in the Northern and Southern United States (20. See many statements in the ‘Ornithological Biography.’ See also some curious observations on the nests of Italian birds by Eugenio Bettoni, in the ‘Atti della Società Italiana,’ vol. xi. 1869, p. 487.), will feel any great difficulty in admitting that birds, either by a change (in the strict sense of the word) of their habits, or through the natural selection of so-called spontaneous variations of instinct, might readily be led to modify their manner of nesting.
This way of viewing the relation, as far as it holds good, between the bright colours of female birds and their manner of nesting, receives some support from certain cases occurring in the Sahara Desert. Here, as in most other deserts, various birds, and many other animals, have had their colours adapted in a wonderful manner to the tints of the surrounding surface. Nevertheless there are, as I am informed by the Rev. Mr. Tristram, some curious exceptions to the rule; thus the male of the Monticola cyanea is conspicuous from his bright blue colour, and the female almost equally conspicuous from her mottled brown and white plumage; both sexes of two species of Dromolaea are of a lustrous black; so that these three species are far from receiving protection from their colours, yet they are able to survive, for they have acquired the habit of taking refuge from danger in holes or crevices in the rocks.
With respect to the above groups in which the females are conspicuously coloured and build concealed nests, it is not necessary to suppose that each separate species had its nidifying instinct specially modified; but only that the early progenitors of each group were gradually led to build domed or concealed nests, and afterwards transmitted this instinct, together with their bright colours, to their modified descendants. As far as it can be trusted, the conclusion is interesting, that sexual selection together with equal or nearly equal inheritance by both sexes, have indirectly determined the manner of nidification of whole groups of birds.
According to Mr. Wallace, even in the groups in which the females, from being protected in domed nests during incubation, have not had their bright colours eliminated through natural selection, the males often differ in a slight, and occasionally in a considerable degree from the females. This is a significant fact, for such differences in colour must be accounted for by some of the variations in the males having been from the first limited in transmission to the same sex; as it can hardly be maintained that these differences, especially when very slight, serve as a protection to the female. Thus all the species in the splendid group of the Trogons build in holes; and Mr. Gould gives figures (21. See his Monograph of the Trogonidae, 1st edition.) of both sexes of twenty-five species, in all of which, with one partial exception, the sexes differ sometimes slightly, sometimes conspicuously, in colour,—the males being always finer than the females, though the latter are likewise beautiful. All the species of kingfishers build in holes, and with most of the species the sexes are equally brilliant, and thus far Mr. Wallace’s rule holds good; but in some of the Australian species the colours of the females are rather less vivid than those of the male; and in one splendidly-coloured species, the sexes differ so much that they were at first thought to be specifically distinct. (22. Namely, Cyanalcyon, Gould’s ‘Handbook to the Birds of Australia,’ vol. i. p. 133; see, also, pp. 130, 136.) Mr. R.B. Sharpe, who has especially studied this group, has shewn me some American species (Ceryle) in which the breast of the male is belted with black. Again, in Carcineutes, the difference between the sexes is conspicuous: in the male the upper surface is dull-blue banded with black, the lower surface being partly fawn-coloured, and there is much red about the head; in the female the upper surface is reddish-brown banded with black, and the lower surface white with black markings. It is an interesting fact, as shewing how the same peculiar style of sexual colouring often characterises allied forms, that in three species of Dacelo the male differs from the female only in the tail being dull-blue banded with black, whilst that of the female is brown with blackish bars; so that here the tail differs in colour in the two sexes in exactly the same manner as the whole upper surface in the two sexes of Carcineutes.
With parrots, which likewise build in holes, we find analogous cases: in most of the species, both sexes are brilliantly coloured and indistinguishable, but in not a few species the males are coloured rather more vividly than the females, or even very differently from them. Thus, besides other strongly-marked differences, the whole under surface of the male King Lory (Aprosmictus scapulatus) is scarlet, whilst the throat and chest of the female is green tinged with red: in the Euphema splendida there is a similar difference, the face and wing coverts moreover of the female being of a paler blue than in the male. (23. Every gradation of difference between the sexes may be followed in the parrots of Australia. See Gould’s ‘Handbook,’ etc., vol. ii. pp. 14-102.) In the family of the tits (Parinae), which build concealed nests, the female of our common blue tomtit (Parus caeruleus), is “much less brightly coloured” than the male: and in the magnificent Sultan yellow tit of India the difference is greater. (24. Macgillivray’s ‘British Birds,’ vol. ii. p. 433. Jerdon, ‘Birds of India,’ vol. ii. p. 282.)