The Descent of Man – Day 118 of 151

Class II

When the adult female is more conspicuous than the adult male, the young of both sexes in their first plumage resemble the adult male.

This class is exactly the reverse of the last, for the females are here brighter coloured or more conspicuous than the males; and the young, as far as they are known, resemble the adult males instead of the adult females. But the difference between the sexes is never nearly so great as with many birds in the first class, and the cases are comparatively rare. Mr. Wallace, who first called attention to the singular relation which exists between the less bright colours of the males and their performing the duties of incubation, lays great stress on this point (13. ‘Westminster Review,’ July 1867, and A. Murray, ‘Journal of Travel,’ 1868, p. 83.), as a crucial test that obscure colours have been acquired for the sake of protection during the period of nesting. A different view seems to me more probable. As the cases are curious and not numerous, I will briefly give all that I have been able to find.

In one section of the genus Turnix, quail-like birds, the female is invariably larger than the male (being nearly twice as large in one of the Australian species), and this is an unusual circumstance with the Gallinaceae. In most of the species the female is more distinctly coloured and brighter than the male (14. For the Australian species, see Gould’s ‘Handbook,’ etc., vol. ii. pp. 178, 180, 186, and 188. In the British Museum specimens of the Australian Plain-wanderer (Pedionomus torquatus) may be seen, shewing similar sexual differences.), but in some few species the sexes are alike. In Turnix taigoor of India the male “wants the black on the throat and neck, and the whole tone of the plumage is lighter and less pronounced than that of the female.” The female appears to be noisier, and is certainly much more pugnacious than the male; so that the females and not the males are often kept by the natives for fighting, like game-cocks. As male birds are exposed by the English bird-catchers for a decoy near a trap, in order to catch other males by exciting their rivalry, so the females of this Turnix are employed in India. When thus exposed the females soon begin their “loud purring call, which can be heard a long way off, and any females within ear-shot run rapidly to the spot, and commence fighting with the caged bird.” In this way from twelve to twenty birds, all breeding females, may be caught in the course of a single day. The natives assert that the females after laying their eggs associate in flocks, and leave the males to sit on them. There is no reason to doubt the truth of this assertion, which is supported by some observations made in China by Mr. Swinhoe. (15. Jerdon, ‘Birds of India,’ vol. iii. p. 596. Mr. Swinhoe, in ‘Ibis,’ 1865, p. 542; 1866, pp. 131, 405.) Mr. Blyth believes, that the young of both sexes resemble the adult male.

Rhynchaea capensis (from Brehm).

Figure 62: Rhynchaea capensis (from Brehm).

The females of the three species of Painted Snipes (Rhynchaea, Fig. 62) “are not only larger but much more richly coloured than the males.” (16. Jerdon, ‘Birds of India,’ vol. iii. p. 677.) With all other birds in which the trachea differs in structure in the two sexes it is more developed and complex in the male than in the female; but in the Rhynchaea australis it is simple in the male, whilst in the female it makes four distinct convolutions before entering the lungs. (17. Gould’s ‘Handbook to the Birds of Australia,’ vol. ii. p. 275.) The female therefore of this species has acquired an eminently masculine character. Mr. Blyth ascertained, by examining many specimens, that the trachea is not convoluted in either sex of R. bengalensis, which species resembles R. australis so closely, that it can hardly be distinguished except by its shorter toes. This fact is another striking instance of the law that secondary sexual characters are often widely different in closely-allied forms, though it is a very rare circumstance when such differences relate to the female sex. The young of both sexes of R. bengalensis in their first plumage are said to resemble the mature male. (18. ‘The Indian Field,’ Sept. 1858, p. 3.) There is also reason to believe that the male undertakes the duty of incubation, for Mr. Swinhoe (19. ‘Ibis,’ 1866, p. 298.) found the females before the close of the summer associated in flocks, as occurs with the females of the Turnix.

The females of Phalaropus fulicarius and P. hyperboreus are larger, and in their summer plumage “more gaily attired than the males.” But the difference in colour between the sexes is far from conspicuous. According to Professor Steenstrup, the male alone of P. fulicarius undertakes the duty of incubation; this is likewise shewn by the state of his breast-feathers during the breeding-season. The female of the dotterel plover (Eudromias morinellus) is larger than the male, and has the red and black tints on the lower surface, the white crescent on the breast, and the stripes over the eyes, more strongly pronounced. The male also takes at least a share in hatching the eggs; but the female likewise attends to the young. (20. For these several statements, see Mr. Gould’s ‘Birds of Great Britain.’ Prof. Newton informs me that he has long been convinced, from his own observations and from those of others, that the males of the above-named species take either the whole or a large share of the duties of incubation, and that they “shew much greater devotion towards their young, when in danger, than do the females.” So it is, as he informs me, with Limosa lapponica and some few other Waders, in which the females are larger and have more strongly contrasted colours than the males.) I have not been able to discover whether with these species the young resemble the adult males more closely than the adult females; for the comparison is somewhat difficult to make on account of the double moult.

Turning now to the ostrich Order: the male of the common cassowary (Casuarius galeatus) would be thought by any one to be the female, from his smaller size and from the appendages and naked skin about his head being much less brightly coloured; and I am informed by Mr. Bartlett that in the Zoological Gardens, it is certainly the male alone who sits on the eggs and takes care of the young. (21. The natives of Ceram (Wallace, ‘Malay Archipelago,’ vol. ii. p. 150) assert that the male and female sit alternately on the eggs; but this assertion, as Mr. Bartlett thinks, may be accounted for by the female visiting the nest to lay her eggs.) The female is said by Mr. T.W. Wood (22. The ‘Student,’ April 1870, p. 124.) to exhibit during the breeding-season a most pugnacious disposition; and her wattles then become enlarged and more brilliantly coloured. So again the female of one of the emus (Dromoeus irroratus) is considerably larger than the male, and she possesses a slight top-knot, but is otherwise indistinguishable in plumage. She appears, however, “to have greater power, when angry or otherwise excited, of erecting, like a turkey-cock, the feathers of her neck and breast. She is usually the more courageous and pugilistic. She makes a deep hollow guttural boom especially at night, sounding like a small gong. The male has a slenderer frame and is more docile, with no voice beyond a suppressed hiss when angry, or a croak.” He not only performs the whole duty of incubation, but has to defend the young from their mother; “for as soon as she catches sight of her progeny she becomes violently agitated, and notwithstanding the resistance of the father appears to use her utmost endeavours to destroy them. For months afterwards it is unsafe to put the parents together, violent quarrels being the inevitable result, in which the female generally comes off conqueror.” (23. See the excellent account of the habits of this bird under confinement, by Mr. A.W. Bennett, in ‘Land and Water,’ May 1868, p. 233.) So that with this emu we have a complete reversal not only of the parental and incubating instincts, but of the usual moral qualities of the two sexes; the females being savage, quarrelsome, and noisy, the males gentle and good. The case is very different with the African ostrich, for the male is somewhat larger than the female and has finer plumes with more strongly contrasted colours; nevertheless he undertakes the whole duty of incubation. (24. Mr. Sclater, on the incubation of the Struthiones, ‘Proc. Zool. Soc.’ June 9, 1863. So it is with the Rhea darwinii: Captain Musters says (‘At Home with the Patagonians,’ 1871, p. 128), that the male is larger, stronger and swifter than the female, and of slightly darker colours; yet he takes sole charge of the eggs and of the young, just as does the male of the common species of Rhea.)

I will specify the few other cases known to me, in which the female is more conspicuously coloured than the male, although nothing is known about the manner of incubation. With the carrion-hawk of the Falkland Islands (Milvago leucurus) I was much surprised to find by dissection that the individuals, which had all their tints strongly pronounced, with the cere and legs orange-coloured, were the adult females; whilst those with duller plumage and grey legs were the males or the young. In an Australian tree-creeper (Climacteris erythrops) the female differs from the male in “being adorned with beautiful, radiated, rufous markings on the throat, the male having this part quite plain.” Lastly, in an Australian night-jar “the female always exceeds the male in size and in the brilliance of her tints; the males, on the other hand, have two white spots on the primaries more conspicuous than in the female.” (25. For the Milvago, see ‘Zoology of the Voyage of the “Beagle,” Birds,’ 1841, p. 16. For the Climacteris and night-jar (Eurostopodus), see Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 602 and 97. The New Zealand shieldrake (Tadorna variegata) offers a quite anomalous case; the head of the female is pure white, and her back is redder than that of the male; the head of the male is of a rich dark bronzed colour, and his back is clothed with finely pencilled slate-coloured feathers, so that altogether he may be considered as the more beautiful of the two. He is larger and more pugnacious than the female, and does not sit on the eggs. So that in all these respects this species comes under our first class of cases; but Mr. Sclater (‘Proceedings of the Zoological Society,’ 1866, p. 150) was much surprised to observe that the young of both sexes, when about three months old, resembled in their dark heads and necks the adult males, instead of the adult females; so that it would appear in this case that the females have been modified, whilst the males and the young have retained a former state of plumage.)

We thus see that the cases in which female birds are more conspicuously coloured than the males, with the young in their immature plumage resembling the adult males instead of the adult females, as in the previous class, are not numerous, though they are distributed in various Orders. The amount of difference, also, between the sexes is incomparably less than that which frequently occurs in the last class; so that the cause of the difference, whatever it may have been, has here acted on the females either less energetically or less persistently than on the males in the last class. Mr. Wallace believes that the males have had their colours rendered less conspicuous for the sake of protection during the period of incubation; but the difference between the sexes in hardly any of the foregoing cases appears sufficiently great for this view to be safely accepted. In some of the cases, the brighter tints of the female are almost confined to the lower surface, and the males, if thus coloured, would not have been exposed to danger whilst sitting on the eggs. It should also be borne in mind that the males are not only in a slight degree less conspicuously coloured than the females, but are smaller and weaker. They have, moreover, not only acquired the maternal instinct of incubation, but are less pugnacious and vociferous than the females, and in one instance have simpler vocal organs. Thus an almost complete transposition of the instincts, habits, disposition, colour, size, and of some points of structure, has been effected between the two sexes.

Now if we might assume that the males in the present class have lost some of that ardour which is usual to their sex, so that they no longer search eagerly for the females; or, if we might assume that the females have become much more numerous than the males–and in the case of one Indian Turnix the females are said to be “much more commonly met with than the males” (26. Jerdon, ‘Birds of India,’ vol. iii. p. 598.)–then it is not improbable that the females would have been led to court the males, instead of being courted by them. This indeed is the case to a certain extent with some birds, as we have seen with the peahen, wild turkey, and certain kinds of grouse. Taking as our guide the habits of most male birds, the greater size and strength as well as the extraordinary pugnacity of the females of the Turnix and emu, must mean that they endeavour to drive away rival females, in order to gain possession of the male; and on this view all the facts become clear; for the males would probably be most charmed or excited by the females which were the most attractive to them by their bright colours, other ornaments, or vocal powers. Sexual selection would then do its work, steadily adding to the attractions of the females; the males and the young being left not at all, or but little modified.

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