The females of the three species of Painted Snipes (Rhynchæa) “are not only larger, but much more richly coloured than the males.”244 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 Rhynchæa Australis it is simple in the male, whilst in the female it makes four distinct convolutions before entering the lungs.245 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 so closely resembles R. Australis 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.246 There is also reason to believe that the male undertakes the duty of incubation, for Mr. Swinhoe247 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. The male alone of P. fulicarius undertakes, according to Professor Steenstrup, the duty of incubation, as 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.248 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.249 The female is said by Mr. T. W. Wood250 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 (Dromœus irroratus) is considerably larger than the male, and she possesses a slight top-knot, but is otherwise undistinguishable 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.”251 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.252
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 their 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 nightjar “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.”253
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 acted on the females in the present class 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 of less size, and have less strength. 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”254—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 and 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 brighter colours, other ornaments, or vocal powers. Sexual selection would then soon 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 both 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 with birds under confinement, in the same manner as often occurs in the case of brilliantly coloured male birds. With many species of herons the young differ greatly from the adults, and their summer plumage, 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 two last classes, on the young having retained a former or ancient state of plumage, which has been exchanged for a new plumage by the old of both sexes. When the adults are brightly 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 two first classes, the transmission, though limited to the same age, has not been limited to the same sex. Consequently both 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 absolutely 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,255—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.256 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.257
Before proceeding, I may remark that under the present and two next 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 some 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 exceeded to a slight degree 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.258 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.259 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 albinoes, as shewn by the colour of their beaks and legs, which nearly resembled the same parts in the adults.260
It may be worth while to illustrate the above three modes by which, in the present class, the two sexes and the young may have come to resemble each other, by the curious case of the genus Passer.261 In the house-sparrow (P. domesticus) the male differs much from the female and from the young. These resemble each other, and likewise to a large extent both sexes and the young of the sparrow of Palestine (P. brachydactylus), as well as of some allied species. We may therefore assume that the female and young of the house-sparrow approximately shew us the plumage of the progenitor of the genus. Now with the tree-sparrow (P. montanus) both sexes and the young closely resemble the male of the house-sparrow; so that they have all been modified in the same manner, and all depart from the typical colouring of their early progenitor. This may have been effected by a male ancestor of the tree-sparrow having varied, firstly, when nearly mature, or, secondly, whilst quite young, having in either case transmitted his modified plumage to the females and the young; or, thirdly, he may have varied when adult and transmitted his plumage to both adult sexes, and, owing to the failure of the law of inheritance at corresponding ages, at some subsequent period to his young.
It is impossible to decide which of these three modes has generally prevailed throughout the present class of cases. The belief that the males varied whilst young, and transmitted their variations to their offspring of both sexes is perhaps the most probable. I may here add that I have endeavoured, with little success, by consulting various works, to decide how far with birds the period of variation has generally determined the transmission of characters to one sex or to both. The two rules, often referred to (namely, that variations occurring late in life are transmitted to one and the same sex, whilst those which occur early in life are transmitted to both sexes), apparently hold good in the first,262 second, and fourth classes of cases; but they fail in an equal number, namely, in the third, often in the fifth,263 and in the sixth small class. They hold good, however, as far as I can judge, with a considerable majority of the species of birds. Whether or not this be so, we may conclude from the facts given in the eighth chapter that the period of variation has been one important element in determining the form of transmission.
With birds it is difficult to decide by what standard we ought to judge of the earliness or lateness of the period of variation, whether by the age in reference to the duration of life, or to the power of reproduction, or to the number of moults through which the species passes. The moulting of birds, even within the same family, sometimes differs much without any assignable cause. Some birds moult so early, that nearly all the body-feathers are cast off before the first wing-feathers are fully grown; and we cannot believe that this was the primordial state of things. When the period of moulting has been accelerated, the age at which the colours of the adult plumage were first developed would falsely appear to us to have been earlier than it really was. This may be illustrated by the practice followed by some bird-fanciers, who pull out a few feathers from the breast of nestling bullfinches, and from the head or neck of young gold pheasants, in order to ascertain their sex; for in the males these feathers are immediately replaced by coloured ones.264 The actual duration of life is known in but few birds, so that we can hardly judge by this standard. And with reference to the period at which the powers of reproduction are gained, it is a remarkable fact that various birds occasionally breed whilst retaining their immature plumage.265
The fact of birds breeding in their immature plumage seems opposed to the belief that sexual selection has played as important a part, as I believe it has, in giving ornamental colours, plumes, &c., to the males, and, by means of equal transmission, to the females of many species. The objection would be a valid one, if the younger and less ornamented males were as successful in winning females and propagating their kind, as the older and more beautiful males. But we have no reason to suppose that this is the case. Audubon speaks of the breeding of the immature males of Ibis tantalus as a rare event, as does Mr. Swinhoe, in regard to the immature males of Oriolus.266 If the young of any species in their immature plumage were more successful in winning partners than the adults, the adult plumage would probably soon be lost, as the males which retained their immature dress for the longest period would prevail, and thus the character of the species would ultimately be modified.267 If, on the other hand, the young never succeeded in obtaining a female, the habit of early reproduction would perhaps be sooner or later quite eliminated, from being superfluous and entailing waste of power.
The plumage of certain birds goes on increasing in beauty during many years after they are fully mature; this is the case with the train of the peacock, and with the crest and plumes of certain herons; for instance, the Ardea Ludovicana;268 but it is very doubtful whether the continued development of such feathers is the result of the selection of successive beneficial variations, or merely of continuous growth. Most fishes continue increasing in size, as long as they are in good health and have plenty of food; and a somewhat similar law may prevail with the plumes of birds.
Class V. When the adults of both sexes have a distinct winter and summer plumage, whether or not the male differs from the female, the young resemble the adults of both sexes in their winter dress, or much more rarely in their summer dress, or they resemble the females alone; or the young may have an intermediate character; or again, they may differ greatly from the adults in both their seasonal plumages.—The cases in this class are singularly complex; nor is this surprising, as they depend on inheritance, limited in a greater or less degree in three different ways, namely by sex, age, and the season of the year. In some cases the individuals of the same species pass through at least five distinct states of plumage. With the species, in which the male differs from the female during the summer season alone, or, which is rarer, during both seasons,269 the young generally resemble the females,—as with the so-called goldfinch of North America, and apparently with the splendid Maluri of Australia.270 With the species, the sexes of which are alike during both the summer and winter, the young may resemble the adults, firstly, in their winter dress; secondly, which occurs much more rarely, in their summer dress; thirdly, they may be intermediate between these two states; and, fourthly, they may differ greatly from the adults at all seasons. We have an instance of the first of these four cases in one of the egrets of India (Buphus coromandus), in which the young and the adults of both sexes are white during the winter, the adults becoming golden-buff during the summer. With the Gaper (Anastomus oscitans) of India we have a similar case, but the colours are reversed; for the young and the adults of both sexes are grey and black during the winter, the adults becoming white during the summer.271 As an instance of the second case, the young of the razor-bill (Alca torda, Linn.), in an early state of plumage, are coloured like the adults during the summer; and the young of the white-crowned sparrow of North America (Fringilla leucophrys), as soon as fledged, have elegant white stripes on their heads, which are lost by the young and the old during the winter.272 With respect to the third case, namely, that of the young having an intermediate character between the summer and winter adult plumages, Yarrell273 insists that this occurs with many waders. Lastly, in regard to the young differing greatly from both sexes in their adult summer and winter plumages, this occurs with some herons and egrets of North America and India,—the young alone being white.
I will make only a few remarks on these complicated cases. When the young resemble the female in her summer dress, or the adults of both sexes in their winter dress, the cases differ from those given under Classes I. and III. only in the characters originally acquired by the males during the breeding-season, having been limited in their transmission to the corresponding season. When the adults have a distinct summer and winter plumage, and the young differ from both, the case is more difficult to understand. We may admit as probable that the young have retained an ancient state of plumage; we can account through sexual selection for the summer or nuptial plumage of the adults, but how are we to account for their distinct winter plumage? If we could admit that this plumage serves in all cases as a protection, its acquirement would be a simple affair; but there seems no good reason for this admission. It may be suggested that the widely different conditions of life during the winter and summer have acted in a direct manner on the plumage; this may have had some effect, but I have not much confidence in so great a difference, as we sometimes see, between the two plumages having been thus caused. A more probable explanation is, that an ancient style of plumage, partially modified through the transference of some characters from the summer plumage, has been retained by the adults during the winter. Finally, all the cases in our present class apparently depend on characters acquired by the adult males, having been variously limited in their transmission according to age, season, and sex; but it would not be worth while to attempt to follow out these complex relations.
Class VI. The young in their first plumage differ from each other according to sex; the young males resembling more or less closely the adult males, and the young females more or less closely the adult females.—The cases in the present class, though occurring in various groups, are not numerous; yet, if experience had not taught us to the contrary, it seems the most natural thing that the young should at first always resemble to a certain extent, and gradually become more and more like, the adults of the same sex. The adult male blackcap (Sylvia atricapilla) has a black head, that of the female being reddish-brown; and I am informed by Mr. Blyth, that the young of both sexes can be distinguished by this character even as nestlings. In the family of thrushes an unusual number of similar cases have been noticed; the male blackbird (Turdus merula) can be distinguished in the nest from the female, as the main wing-feathers, which are not moulted so soon as the body-feathers, retain a brownish tint until the second general moult.274 The two sexes of the mocking bird (Turdus polyglottus, Linn.) differ very little from each other, yet the males can easily be distinguished at a very early age from the females by shewing more pure white.275 The males of a forest-thrush and of a rock-thrush (viz. Orocetes erythrogastra and Petrocincla cyanea) have much of their plumage of a fine blue, whilst the females are brown; and the nestling males of both species have their main wing and tail-feathers edged with blue, whilst those of the female are edged with brown.276 So that the very same feathers which in the young blackbird assume their mature character and become black after the others, in these two species assume this character and become blue before the others. The most probable view with reference to these cases is that the males, differently from what occurs in Class I., have transmitted their colours to their male offspring at an earlier age than that at which they themselves first acquired them; for if they had varied whilst quite young, they would probably have transmitted all their characters to their offspring of both sexes.277
In Aïthurus polytmus (one of the humming-birds) the male is splendidly coloured black and green, and two of the tail-feathers are immensely lengthened; the female has an ordinary tail and inconspicuous colours; now the young males, instead of resembling the adult female, in accordance with the common rule, begin from the first to assume the colours proper to their sex, and their tail-feathers soon become elongated. I owe this information to Mr. Gould, who has given me the following more striking and as yet unpublished case. Two humming-birds belonging to the genus Eustephanus, both beautifully coloured, inhabit the small island of Juan Fernandez, and have always been ranked as specifically distinct. But it has lately been ascertained that the one, which is of a rich chesnut-brown colour with a golden-red head, is the male, whilst the other, which is elegantly variegated with green and white with a metallic-green head, is the female. Now the young from the first resemble to a certain extent the adults of the corresponding sex, the resemblance gradually becoming more and more complete.
In considering this last case, if as before we take the plumage of the young as our guide, it would appear that both sexes have been independently rendered beautiful; and not that the one sex has partially transferred its beauty to the other. The male apparently has acquired his bright colours through sexual selection in the same manner as, for instance, the peacock or pheasant in our first class of cases; and the female in the same manner as the female Rhynchæa or Turnix in our second class of cases. But there is much difficulty in understanding how this could have been effected at the same time with the two sexes of the same species. Mr. Salvin states, as we have seen in the eighth chapter, that with certain humming-birds the males greatly exceed in number the females, whilst with other species inhabiting the same country the females greatly exceed the males. If, then, we might assume that during some former lengthened period the males of the Juan Fernandez species had greatly exceeded the females in number, but that during another lengthened period the females had greatly exceeded the males, we could understand how the males at one time, and the females at another time, might have been rendered beautiful by the selection of the brighter-coloured individuals of either sex; both sexes transmitting their characters to their young at a rather earlier age than usual. Whether this is the true explanation I will not pretend to say; but the case is too remarkable to be passed over without notice.
We have now seen in numerous instances under all six classes, that an intimate relation exists between the plumage of the young and that of the adults, either of one sex or both sexes. These relations are fairly well explained on the principle that one sex—this being in the great majority of cases the male—first acquired through variation and sexual selection bright colours or other ornaments, and transmitted them in various ways, in accordance with the recognised laws of inheritance. Why variations have occurred at different periods of life, even sometimes with the species of the same group, we do not know; but with respect to the form of transmission, one important determining cause seems to have been the age at which the variations first appeared.
From the principle of inheritance at corresponding ages, and from any variations in colour which occurred in the males at an early age not being then selected, on the contrary being often eliminated as dangerous, whilst similar variations occurring at or near the period of reproduction have been preserved, it follows that the plumage of the young will often have been left unmodified, or but little modified. We thus get some insight into the colouring of the progenitors of our existing species. In a vast number of species in five out of our six classes of cases, the adults of one sex or both are brightly coloured, at least during the breeding-season, whilst the young are invariably less brightly coloured than the adults, or are quite dull-coloured; for no instance is known, as far as I can discover, of the young of dull-coloured species displaying bright colours, or of the young of brightly-coloured species being more brilliantly coloured than their parents. In the fourth class, however, in which the young and the old resemble each other, there are many species (though by no means all) brightly-coloured, and as these form whole groups, we may infer that their early progenitors were likewise brightly-coloured. With this exception, if we look to the birds of the world, it appears that their beauty has been greatly increased since that period, of which we have a partial record in their immature plumage.
On the Colour of the Plumage in relation to Protection.—It will have been seen that I cannot follow Mr. Wallace in the belief that dull colours when confined to the females have been in most cases specially gained for the sake of protection. There can, however, be no doubt, as formerly remarked, that both sexes of many birds have had their colours modified for this purpose, so as to escape the notice of their enemies; or, in some instances, so as to approach their prey unobserved, in the same manner as owls have had their plumage rendered soft, that their flight may not be overheard. Mr. Wallace remarks278 that “it is only in the tropics, among forests which never lose their foliage, that we find whole groups of birds, whose chief colour is green.” It will be admitted by every one, who has ever tried, how difficult it is to distinguish parrots in a leaf-covered tree. Nevertheless, we must remember that many parrots are ornamented with crimson, blue, and orange tints, which can hardly be protective. Woodpeckers are eminently arboreal, but, besides green species, there are many black, and black-and-white kinds—all the species being apparently exposed to nearly the same dangers. It is therefore probable that strongly-pronounced colours have been acquired by tree-haunting birds through sexual selection, but that green tints have had an advantage through natural selection over other colours for the sake of protection.
In regard to birds which live on the ground, everyone admits that they are coloured so as to imitate the surrounding surface. How difficult it is to see a partridge, snipe, woodcock, certain plovers, larks, and nightjars when crouched on the ground. Animals inhabiting deserts offer the most striking instances, for the bare surface affords no concealment, and all the smaller quadrupeds, reptiles, and birds depend for safety on their colours. As Mr. Tristram has remarked,279 in regard to the inhabitants of the Sahara, all are protected by their “isabelline or sand-colour.” Calling to my recollection the desert-birds which I had seen in South America, as well as most of the ground-birds in Great Britain, it appeared to me that both sexes in such cases are generally coloured nearly alike. Accordingly I applied to Mr. Tristram, with respect to the birds of the Sahara, and he has kindly given me the following information. There are twenty-six species, belonging to fifteen genera, which manifestly have had their plumage coloured in a protective manner; and this colouring is all the more striking, as with most of these birds it is different from that of their congeners. Both sexes of thirteen out of the twenty-six species are coloured in the same manner; but these belong to genera in which this rule commonly prevails, so that they tell us nothing about the protective colours being the same in both sexes of desert-birds. Of the other thirteen species, three belong to genera in which the sexes usually differ from each other, yet they have the sexes alike. In the remaining ten species, the male differs from the female; but the difference is confined chiefly to the under surface of the plumage, which is concealed when the bird crouches on the ground; the head and back being of the same sand-coloured hue in both sexes. So that in these ten species the upper surfaces of both sexes have been acted on and rendered alike, through natural selection, for the sake of protection; whilst the lower surfaces of the males alone have been diversified through sexual selection, for the sake of ornament. Here, as both sexes are equally well protected, we clearly see that the females have not been prevented through natural selection from inheriting the colours of their male parents: we must look to the law of sexually limited transmission, as before explained.
In all parts of the world both sexes of many soft-billed birds, especially those which frequent reeds or sedges, are obscurely coloured. No doubt if their colours had been brilliant, they would have been much more conspicuous to their enemies; but whether their dull tints have been specially gained for the sake of protection seems, as far as I can judge, rather doubtful. It is still more doubtful whether such dull tints can have been gained for the sake of ornament. We must, however, bear in mind that male birds, though dull-coloured, often differ much from their females, as with the common sparrow, and this leads to the belief that such colours have been gained through sexual selection, from being attractive. Many of the soft-billed birds are songsters; and a discussion in a former chapter should not be forgotten, in which it was shewn that the best songsters are rarely ornamented with bright tints. It would appear that female birds, as a general rule, have selected their mates either for their sweet voices or gay colours, but not for both charms combined. Some species which are manifestly coloured for the sake of protection, such as the jack-snipe, woodcock, and nightjar, are likewise marked and shaded, according to our standard of taste, with extreme elegance. In such cases we may conclude that both natural and sexual selection have acted conjointly for protection and ornament. Whether any bird exists which does not possess some special attraction, by which to charm the opposite sex, may be doubted. When both sexes are so obscurely coloured, that it would be rash to assume the agency of sexual selection, and when no direct evidence can be advanced shewing that such colours serve as a protection, it is best to own complete ignorance of the cause, or, which comes to nearly the same thing, to attribute the result to the direct action of the conditions of life.
There are many birds both sexes of which are conspicuously, though not brilliantly coloured, such as the numerous black, white, or piebald species; and these colours, are probably the result of sexual selection. With the common blackbird, capercailzie, black-cock, black Scoter-duck (Oidemia), and even with one of the Birds of Paradise (Lophorina atra), the males alone are black, whilst the females are brown or mottled; and there can hardly be a doubt that blackness in these cases has been a sexually selected character. Therefore it is in some degree probable that the complete or partial blackness of both sexes in such birds as crows, certain cockatoos, storks, and swans, and many marine birds, is likewise the result of sexual selection, accompanied by equal transmission to both sexes; for blackness can hardly serve in any case as a protection. With several birds, in which the male alone is black, and in others in which both sexes are black, the beak or skin about the head is brightly coloured, and the contrast thus afforded adds greatly to their beauty; we see this in the bright yellow beak of the male blackbird, in the crimson skin over the eyes of the black-cock and capercailzie, in the variously and brightly-coloured beak of the Scoter-drake (Oidemia), in the red beak of the chough (Corvus graculus, Linn.), of the black swan, and black stork. This leads me to remark that it is not at all incredible that toucans may owe the enormous size of their beaks to sexual selection, for the sake of displaying the diversified and vivid stripes of colour, with which these organs are ornamented.280 The naked skin at the base of the beak and round the eyes is likewise often brilliantly coloured; and Mr. Gould, in speaking of one species,281 says that the colours of the beak “are doubtless in the finest and most brilliant state during the time of pairing.” There is no greater improbability in toucans being encumbered with immense beaks, though rendered as light as possible by their cancellated structure, for an object falsely appearing to us unimportant, namely, the display of fine colours, than that the male Argus pheasant and some other birds should be encumbered with plumes so long as to impede their flight.
In the same manner, as the males alone of various species are black, the females being dull-coloured; so in a few cases the males alone are either wholly or partially white, as with the several Bell-birds of South America (Chasmorhynchus), the Antarctic goose (Bernicla antarctica), the silver pheasant, &c., whilst the females are brown or obscurely mottled. Therefore, on the same principle as before, it is probable that both sexes of many birds, such as white cockatoos, several egrets with their beautiful plumes, certain ibises, gulls, terns, &c., have acquired their more or less completely white plumage through sexual selection. The species which inhabit snowy regions of course come under a different head. The white plumage of some of the above-named birds appears in both sexes only when they are mature. This is likewise the case with certain gannets, tropic-birds, &c., and with the snow-goose (Anser hyperboreus). As the latter breeds on the “barren grounds,” when not covered with snow, and as it migrates southward during the winter, there is no reason to suppose that its snow-white adult plumage serves as a protection. In the case of the Anastomus oscitans previously alluded to, we have still better evidence that the white plumage is a nuptial character, for it is developed only during the summer; the young in their immature state, and the adults in their winter dress, being grey and black. With many kinds of gulls (Larus), the head and neck become pure white during the summer, being grey or mottled during the winter and in the young state. On the other hand, with the smaller gulls, or sea-mews (Gavia), and with some terns (Sterna), exactly the reverse occurs; for the heads of the young birds during the first year, and of the adults during the winter, are either pure white, or much paler-coloured than during the breeding-season. These latter cases offer another instance of the capricious manner in which sexual selection appears often to have acted.282
The cause of aquatic birds having acquired a white plumage so much more frequently than terrestrial birds, probably depends on their large size and strong powers of flight, so that they can easily defend themselves or escape from birds of prey, to which moreover they are not much exposed. Consequently sexual selection has not here been interfered with or guided for the sake of protection. No doubt, with birds which roam over the open ocean, the males and females could find each other much more easily when made conspicuous either by being perfectly white, or intensely black; so that these colours may possibly serve the same end as the call-notes of many land-birds. A white or black bird, when it discovers and flies down to a carcase floating on the sea or cast up on the beach, will be seen from a great distance, and will guide other birds of the same and of distinct species, to the prey; but as this would be a disadvantage to the first finders, the individuals which were the whitest or blackest would not thus have procured more food than the less strongly coloured individuals. Hence conspicuous colours cannot have been gradually acquired for this purpose through natural selection.283
As sexual selection depends on so fluctuating an element as taste, we can understand how it is that within the same group of birds, with habits of life nearly the same, there should exist white or nearly white, as well as black, or nearly black species,—for instance, white and black cockatoos, storks, ibises, swans, terns, and petrels. Piebald birds likewise sometimes occur in the same groups, for instance, the black-necked swan, certain terns, and the common magpie. That a strong contrast in colour is agreeable to birds, we may conclude, by looking through any large collection of specimens or series of coloured plates, for the sexes frequently differ from each other in the male having the pale parts of a purer white, and the variously coloured dark parts of still darker tints than in the female.
It would even appear that mere novelty, or change for the sake of change, has sometimes acted like a charm on female birds, in the same manner as changes of fashion with us. The Duke of Argyll says,284—and I am glad to have the unusual satisfaction of following for even a short distance in his footsteps—“I am more and more convinced that variety, mere variety, must be admitted to be an object and an aim in Nature.” I wish the Duke had explained what he here means by Nature. Is it meant that the Creator of the universe ordained diversified results for His own satisfaction, or for that of man? The former notion seems to me as much wanting in due reverence as the latter in probability. Capriciousness of taste in the birds themselves appears a more fitting explanation. For example; the males of some parrots can hardly be said to be more beautiful, at least according to our taste, than the females, but they differ from them in such points, as the male having a rose-coloured collar instead of, as in the female, “a bright emeraldine narrow green collar;” or in the male having a black collar instead of “a yellow demi-collar in front,” with a pale roseate instead of a plum-blue head.285 As so many male birds have for their chief ornament elongated tail-feathers or elongated crests, the shortened tail, formerly described in the male of a humming-bird, and the shortened crest of the male goosander almost seem like one of the many opposite changes of fashion which we admire in our own dresses.
Some members of the heron family offer a still more curious case of novelty in colouring having apparently been appreciated for the sake of novelty. The young of the Ardea asha are white, the adults being dark slate-coloured; and not only the young, but the adults of the allied Buphus coromandus in their winter plumage are white, this colour changing into a rich golden-buff during the breeding-season. It is incredible that the young of these two species, as well as of some other members of the same family,286 should have been specially rendered pure white and thus made conspicuous to their enemies; or that the adults of one of these two species should have been specially rendered white during the winter in a country which is never covered with snow. On the other hand we have reason to believe that whiteness has been gained by many birds as a sexual ornament. We may therefore conclude that an early progenitor of the Ardea asha and the Buphus acquired a white plumage for nuptial purposes, and transmitted this colour to their young; so that the young and the old became white like certain existing egrets; the whiteness having afterwards been retained by the young whilst exchanged by the adults for more strongly pronounced tints. But if we could look still further backwards in time to the still earlier progenitors of these two species, we should probably see the adults dark-coloured. I infer that this would be the case, from the analogy of many other birds, which are dark whilst young, and when adult are white; and more especially from the case of the Ardea gularis, the colours of which are the reverse of those of A. asha, for the young are dark-coloured and the adults white, the young having retained a former state of plumage. It appears therefore that the progenitors in their adult condition of the Ardea asha, the Buphus, and of some allies, have undergone, during a long line of descent, the following changes of colour: firstly a dark shade, secondly pure white, and thirdly, owing to another change of fashion (if I may so express myself), their present slaty, reddish, or golden-buff tints. These successive changes are intelligible only on the principle of novelty having been admired by birds for the sake of novelty.
Summary of the Four Chapters on Birds.—Most male birds are highly pugnacious during the breeding-season, and some possess weapons especially adapted for fighting with their rivals. But the most pugnacious and the best-armed males rarely or never depend for success solely on their power to drive away or kill their rivals, but have special means for charming the female. With some it is the power of song, or of emitting strange cries, or of producing instrumental music, and the males in consequence differ from the females in their vocal organs, or in the structure of certain feathers. From the curiously diversified means for producing various sounds we gain a high idea of the importance of this means of courtship. Many birds endeavour to charm the females by love-dances or antics, performed on the ground or in the air, and sometimes at prepared places. But ornaments of many kinds, the most brilliant tints, combs and wattles, beautiful plumes, elongated feathers, top-knots, and so forth, are by far the commonest means. In some cases mere novelty appears to have acted as a charm. The ornaments of the males must be highly important to them, for they have been acquired in not a few cases at the cost of increased danger from enemies, and even at some loss of power in fighting with their rivals. The males of very many species do not assume their ornamental dress until they arrive at maturity, or they assume it only during the breeding-season, or the tints then become more vivid. Certain ornamental appendages become enlarged, turgid, and brightly-coloured during the very act of courtship. The males display their charms with elaborate care and to the best effect; and this is done in the presence of the females. The courtship is sometimes a prolonged affair, and many males and females congregate at an appointed place. To suppose that the females do not appreciate the beauty of the males is to admit that their splendid decorations, all their pomp and display, are useless; and this is incredible. Birds have fine powers of discrimination, and in some few instances it can be shewn that they have a taste for the beautiful. The females, moreover, are known occasionally to exhibit a marked preference or antipathy for certain individual males.
If it be admitted that the females prefer, or are unconsciously excited by the more beautiful males, then the males would slowly but surely be rendered more and more attractive through sexual selection. That it is this sex which has been chiefly modified we may infer from the fact that in almost every genus in which the sexes differ, the males differ much more from each other than do the females; this is well shewn in certain closely-allied representative species in which the females can hardly be distinguished, whilst the males are quite distinct. Birds in a state of nature offer individual differences which would amply suffice for the work of sexual selection; but we have seen that they occasionally present more strongly-marked variations which recur so frequently that they would immediately be fixed, if they served to allure the female. The laws of variation will have determined the nature of the initial changes, and largely influenced the final result. The gradations, which may be observed between the males of allied species, indicate the nature of the steps which have been passed through, and explain in the most interesting manner certain characters, such as the indented ocelli of the tail-feathers of the peacock, and the wonderfully-shaded ocelli of the wing-feathers of the Argus pheasant. It is evident that the brilliant colours, top-knots, fine plumes, &c., of many male birds cannot have been acquired as a protection; indeed they sometimes lead to danger. That they are not due to the direct and definite action of the conditions of life, we may feel assured, because the females have been exposed to the same conditions, and yet often differ from the males to an extreme degree. Although it is probable that changed conditions acting during a lengthened period have produced some definite effect on both sexes, the more important result will have been an increased tendency to fluctuating variability or to augmented individual differences; and such differences will have afforded an excellent groundwork for the action of sexual selection.
The laws of inheritance, irrespectively of selection, appear to have determined whether the characters acquired by the males for the sake of ornament, for producing various sounds, and for fighting together, have been transmitted to the males alone or to both sexes, either permanently or periodically during certain seasons of the year. Why various characters should sometimes have been transmitted in one way and sometimes in another is, in most cases, not known; but the period of variability seems often to have been the determining cause. When the two sexes have inherited all characters in common they necessarily resemble each other; but as the successive variations may be differently transmitted, every possible gradation may be found, even within the same genus, from the closest similarity to the widest dissimilarity between the sexes. With many closely-allied species, following nearly the same habits of life, the males have come to differ from each other chiefly through the action of sexual selection; whilst the females have come to differ chiefly from partaking in a greater or lesser degree of the characters thus acquired by the males. The effects, moreover, of the definite action of the conditions of life, will not have been masked in the females, as in the case of the males, by the accumulation through sexual selection of strongly-pronounced colours and other ornaments. The individuals of both sexes, however affected, will have been kept at each successive period nearly uniform by the free intercrossing of many individuals.
With the species, in which the sexes differ in colour, it is possible that at first there existed a tendency to transmit the successive variations equally to both sexes; and that the females were prevented from acquiring the bright colours of the males, on account of the danger to which they would have been exposed during incubation. But it would be, as far as I can see, an extremely difficult process to convert, by means of natural selection, one form of transmission into another. On the other hand there would not be the least difficulty in rendering a female dull-coloured, the male being still kept bright-coloured, by the selection of successive variations, which were from the first limited in their transmission to the same sex. Whether the females of many species have actually been thus modified, must at present remain doubtful. When, through the law of the equal transmission of characters to both sexes, the females have been rendered as conspicuously coloured as the males, their instincts have often been modified, and they have been led to build domed or concealed nests.
In one small and curious class of cases the characters and habits of the two sexes have been completely transposed, for the females are larger, stronger, more vociferous and brightly-coloured than their males. They have, also, become so quarrelsome that they often fight together like the males of the most pugnacious species. If, as seems probable, they habitually drive away rival females, and by the display of their bright colours or other charms endeavour to attract the males, we can understand how it is that they have gradually been rendered, by means of sexual selection and sexually-limited transmission, more beautiful than the males—the latter being left unmodified or only slightly modified.
Whenever the law of inheritance at corresponding ages prevails, but not that of sexually-limited transmission, then if the parents vary late in life—and we know that this constantly occurs with our poultry, and occasionally with other birds—the young will be left unaffected, whilst the adults of both sexes will be modified. If both these laws of inheritance prevail and either sex varies late in life, that sex alone will be modified, the other sex and the young being left unaffected. When variations in brightness or in other conspicuous characters occur early in life, as no doubt often happens, they will not be acted on through sexual selection until the period of reproduction arrives; consequently they will be liable to be lost by the accidental deaths of the young, and if dangerous will be eliminated through natural selection. Thus we can understand how it is that variations arising late in life have chiefly been preserved for the ornamentation and arming of the males, the females and the young being left almost unaffected, and therefore like each other. With species having a distinct summer and winter plumage, the males of which either resemble or differ from the females during both seasons or during the summer alone, the degrees and kinds of resemblance between the young and the old are exceedingly complex; and this complexity apparently depends on characters, first acquired by the males, being transmitted in various ways and degrees, as limited by age, sex, and season.
As the young of so many species have been but little modified in colour and in other ornaments, we are enabled to form some judgment with respect to the plumage of their early progenitors; and we may infer that the beauty of our existing species, if we look to the whole class, has been largely increased since that period of which the immature plumage gives us an indirect record. Many birds, especially those which live much on the ground, have undoubtedly been obscurely coloured for the sake of protection. In some instances the upper exposed surface of the plumage has been thus coloured in both sexes, whilst the lower surface in the males alone has been variously ornamented through sexual selection. Finally, from the facts given in these four chapters, we may conclude that weapons for battle, organs for producing sound, ornaments of many kinds, bright and conspicuous colours, have generally been acquired by the males through variation and sexual selection, and have been transmitted in various ways according to the several laws of inheritance—the females and the young being left comparatively but little modified.287