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Descent of Man and Selection in Relation to Sex, Part 2

No one doubts that both sexes of many birds have had their colours adapted for the sake of protection; and it is possible that the females alone of some species may have been modified for this end. Although it would be a difficult, perhaps an impossible process, as shewn in the last chapter, to convert one form of transmission into another through selection, there would not be the least difficulty in adapting the colours of the female, independently of those of the male, to surrounding objects, through the accumulation of variations which were from the first limited in their transmission to the female sex. If the variations were not thus limited, the bright tints of the male would be deteriorated or destroyed. Whether the females alone of many species have been thus specially modified, is at present very doubtful. I wish I could follow Mr. Wallace to the full extent; for the admission would remove some difficulties. Any variations which were of no service to the female as a protection would be at once obliterated, instead of being lost simply by not being selected, or from free intercrossing, or from being eliminated when transferred to the male and in any way injurious to him. Thus the plumage of the female would be kept constant in character. It would also be a relief if we could admit that the obscure tints of both sexes of many birds had been acquired and preserved for the sake of protection,—for example, of the hedge-warbler or kitty-wren (Accentor modularis and Troglodytes vulgaris), with respect to which we have no sufficient evidence of the action of sexual selection. We ought, however, to be cautious in concluding that colours which appear to us dull, are not attractive to the females of certain species; we should bear in mind such cases as that of the common house-sparrow, in which the male differs much from the female, but does not exhibit any bright tints. No one probably will dispute that many gallinaceous birds which live on the open ground, have acquired their present colours, at least in part, for the sake of protection. We know how well they are thus concealed; we know that ptarmigans, whilst changing from their winter to their summer plumage, both of which are protective, suffer greatly from birds of prey. But can we believe that the very slight differences in tints and markings between, for instance, the female black-grouse and red-grouse serve as a protection? Are partridges, as they are now coloured, better protected than if they had resembled quails? Do the slight differences between the females of the common pheasant, the Japan and gold pheasants, serve as a protection, or might not their plumages have been interchanged with impunity? From what Mr. Wallace has observed of the habits of certain gallinaceous birds in the East, he thinks that such slight differences are beneficial. For myself, I will only say that I am not convinced. Formerly when I was inclined to lay much stress on protection as accounting for the duller colours of female birds, it occurred to me that possibly both sexes and the young might aboriginally have been equally bright coloured; but that subsequently, the females from the danger incurred during incubation, and the young from being inexperienced, had been rendered dull as a protection. But this view is not supported by any evidence, and is not probable; for we thus in imagination expose during past times the females and the young to danger, from which it has subsequently been necessary to shield their modified descendants. We have, also, to reduce, through a gradual process of selection, the females and the young to almost exactly the same tints and markings, and to transmit them to the corresponding sex and period of life. On the supposition that the females and the young have partaken during each stage of the process of modification of a tendency to be as brightly coloured as the males, it is also a somewhat strange fact that the females have never been rendered dull-coloured without the young participating in the same change; for there are no instances, as far as I can discover, of species with the females dull and the young bright coloured. A partial exception, however, is offered by the young of certain woodpeckers, for they have “the whole upper part of the head tinged with red,” which afterwards either decreases into a mere circular red line in the adults of both sexes, or quite disappears in the adult females. (12. Audubon, ‘Ornith. Biography,’ vol. i. p. 193. Macgillivray, ‘History of British Birds,’ vol. iii. p. 85. See also the case before given of Indopicus carlotta.) Finally, with respect to our present class of cases, the most probable view appears to be that successive variations in brightness or in other ornamental characters, occurring in the males at a rather late period of life have alone been preserved; and that most or all of these variations, owing to the late period of life at which they appeared, have been from the first transmitted only to the adult male offspring. Any variations in brightness occurring in the females or in the young, would have been of no service to them, and would not have been selected; and moreover, if dangerous, would have been eliminated. Thus the females and the young will either have been left unmodified, or (as is much more common) will have been partially modified by receiving through transference from the males some of his successive variations. Both sexes have perhaps been directly acted on by the conditions of life to which they have long been exposed: but the females from not being otherwise much modified, will best exhibit any such effects. These changes and all others will have been kept uniform by the free intercrossing of many individuals. In some cases, especially with ground birds, the females and the young may possibly have been modified, independently of the males, for the sake of protection, so as to have acquired the same dull-coloured plumage. 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. [Fig. 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. CLASS III. — WHEN THE ADULT MALE RESEMBLES THE ADULT FEMALE, THE YOUNG OF BOTH SEXES HAVE A PECULIAR FIRST PLUMAGE OF THEIR OWN. In this class the sexes when adult resemble each other, and differ from the young. This occurs with many birds of many kinds. The male robin can hardly be distinguished from the female, but the young are widely different, with their mottled dusky-olive and brown plumage. The male and female of the splendid scarlet ibis are alike, whilst the young are brown; and the scarlet colour, though common to both sexes, is apparently a sexual character, for it is not well developed in either sex under confinement; and a loss of colour often occurs with brilliant males when they are confined. With many species of herons the young differ greatly from the adults; and the summer plumage of the latter, though common to both sexes, clearly has a nuptial character. Young swans are slate-coloured, whilst the mature birds are pure white; but it would be superfluous to give additional instances. These differences between the young and the old apparently depend, as in the last two classes, on the young having retained a former or ancient state of plumage, whilst the old of both sexes have acquired a new one. When the adults are bright coloured, we may conclude from the remarks just made in relation to the scarlet ibis and to many herons, and from the analogy of the species in the first class, that such colours have been acquired through sexual selection by the nearly mature males; but that, differently from what occurs in the first two classes, the transmission, though limited to the same age, has not been limited to the same sex. Consequently, the sexes when mature resemble each other and differ from the young. CLASS IV. — WHEN THE ADULT MALE RESEMBLES THE ADULT FEMALE, THE YOUNG OF BOTH SEXES IN THEIR FIRST PLUMAGE RESEMBLE THE ADULTS. In this class the young and the adults of both sexes, whether brilliantly or obscurely coloured, resemble each other. Such cases are, I think, more common than those in the last class. We have in England instances in the kingfisher, some woodpeckers, the jay, magpie, crow, and many small dull-coloured birds, such as the hedge-warbler or kitty-wren. But the similarity in plumage between the young and the old is never complete, and graduates away into dissimilarity. Thus the young of some members of the kingfisher family are not only less vividly coloured than the adults, but many of the feathers on the lower surface are edged with brown (27. Jerdon, ‘Birds of India,’ vol. i. pp. 222, 228. Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 124, 130.),—a vestige probably of a former state of the plumage. Frequently in the same group of birds, even within the same genus, for instance in an Australian genus of parrakeets (Platycercus), the young of some species closely resemble, whilst the young of other species differ considerably, from their parents of both sexes, which are alike. (28. Gould, ibid. vol. ii. pp. 37, 46, 56.) Both sexes and the young of the common jay are closely similar; but in the Canada jay (Perisoreus canadensis) the young differ so much from their parents that they were formerly described as distinct species. (29. Audubon, ‘Ornith. Biography,’ vol. ii. p. 55.) I may remark before proceeding that, under the present and next two classes of cases, the facts are so complex and the conclusions so doubtful, that any one who feels no especial interest in the subject had better pass them over. The brilliant or conspicuous colours which characterise many birds in the present class, can rarely or never be of service to them as a protection; so that they have probably been gained by the males through sexual selection, and then transferred to the females and the young. It is, however, possible that the males may have selected the more attractive females; and if these transmitted their characters to their offspring of both sexes, the same results would follow as from the selection of the more attractive males by the females. But there is evidence that this contingency has rarely, if ever, occurred in any of those groups of birds in which the sexes are generally alike; for, if even a few of the successive variations had failed to be transmitted to both sexes, the females would have slightly exceeded the males in beauty. Exactly the reverse occurs under nature; for, in almost every large group in which the sexes generally resemble each other, the males of some few species are in a slight degree more brightly coloured than the females. It is again possible that the females may have selected the more beautiful males, these males having reciprocally selected the more beautiful females; but it is doubtful whether this double process of selection would be likely to occur, owing to the greater eagerness of one sex than the other, and whether it would be more efficient than selection on one side alone. It is, therefore, the most probable view that sexual selection has acted, in the present class, as far as ornamental characters are concerned, in accordance with the general rule throughout the animal kingdom, that is, on the males; and that these have transmitted their gradually-acquired colours, either equally or almost equally, to their offspring of both sexes. Another point is more doubtful, namely, whether the successive variations first appeared in the males after they had become nearly mature, or whilst quite young. In either case sexual selection must have acted on the male when he had to compete with rivals for the possession of the female; and in both cases the characters thus acquired have been transmitted to both sexes and all ages. But these characters if acquired by the males when adult, may have been transmitted at first to the adults alone, and at some subsequent period transferred to the young. For it is known that, when the law of inheritance at corresponding ages fails, the offspring often inherit characters at an earlier age than that at which they first appeared in their parents. (30. ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 79.) Cases apparently of this kind have been observed with birds in a state of nature. For instance Mr. Blyth has seen specimens of Lanius rufus and of Colymbus glacialis which had assumed whilst young, in a quite anomalous manner, the adult plumage of their parents. (31. ‘Charlesworth’s Magazine of Natural History,’ vol. i. 1837, pp. 305, 306.) Again, the young of the common swan (Cygnus olor) do not cast off their dark feathers and become white until eighteen months or two years old; but Dr. F. Forel has described the case of three vigorous young birds, out of a brood of four, which were born pure white. These young birds were not albinos, as shewn by the colour of their beaks and legs, which nearly resembled the same parts in the adults. (32. ‘Bulletin de la Soc. Vaudoise des Sc. Nat.’ vol. x. 1869, p. 132. The young of the Polish swan, Cygnus immutabilis of Yarrell, are always white; but this species, as Mr. Sclater informs me, is believed to be nothing more than a variety of the domestic swan (Cygnus olor).) <div style="break-after:column;"></div>

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