XV. BIRDS—continued…

XV. BIRDS—continued

Discussion as to why the males alone of some species, and both sexes of others, are brightly coloured—On sexually-limited inheritance, as applied to various structures and to brightly-coloured plumage—Nidification in relation to colour—Loss of nuptial plumage during the winter.

We have in this chapter to consider why the females of many birds have not acquired the same ornaments as the male; and why, on the other hand, both sexes of many other birds are equally, or almost equally, ornamented? In the following chapter we shall consider the few cases in which the female is more conspicuously coloured than the male.

In my ‘Origin of Species’ (1. Fourth edition, 1866, p. 241.) I briefly suggested that the long tail of the peacock would be inconvenient and the conspicuous black colour of the male capercailzie dangerous, to the female during the period of incubation: and consequently that the transmission of these characters from the male to the female offspring had been checked through natural selection. I still think that this may have occurred in some few instances: but after mature reflection on all the facts which I have been able to collect, I am now inclined to believe that when the sexes differ, the successive variations have generally been from the first limited in their transmission to the same sex in which they first arose. Since my remarks appeared, the subject of sexual coloration has been discussed in some very interesting papers by Mr. Wallace (2. ‘Westminster Review,’ July 1867. ‘Journal of Travel,’ vol. i. 1868, p. 73.), who believes that in almost all cases the successive variations tended at first to be transmitted equally to both sexes; but that the female was saved, through natural selection, from acquiring the conspicuous colours of the male, owing to the danger which she would thus have incurred during incubation.

This view necessitates a tedious discussion on a difficult point, namely, whether the transmission of a character, which is at first inherited by both sexes can be subsequently limited in its transmission to one sex alone by means of natural selection. We must bear in mind, as shewn in the preliminary chapter on sexual selection, that characters which are limited in their development to one sex are always latent in the other. An imaginary illustration will best aid us in seeing the difficulty of the case; we may suppose that a fancier wished to make a breed of pigeons, in which the males alone should be coloured of a pale blue, whilst the females retained their former slaty tint. As with pigeons characters of all kinds are usually transmitted to both sexes equally, the fancier would have to try to convert this latter form of inheritance into sexually-limited transmission. All that he could do would be to persevere in selecting every male pigeon which was in the least degree of a paler blue; and the natural result of this process, if steadily carried on for a long time, and if the pale variations were strongly inherited or often recurred, would be to make his whole stock of a lighter blue. But our fancier would be compelled to match, generation after generation, his pale blue males with slaty females, for he wishes to keep the latter of this colour. The result would generally be the production either of a mongrel piebald lot, or more probably the speedy and complete loss of the pale-blue tint; for the primordial slaty colour would be transmitted with prepotent force. Supposing, however, that some pale-blue males and slaty females were produced during each successive generation, and were always crossed together, then the slaty females would have, if I may use the expression, much blue blood in their veins, for their fathers, grandfathers, etc., will all have been blue birds. Under these circumstances it is conceivable (though I know of no distinct facts rendering it probable) that the slaty females might acquire so strong a latent tendency to pale-blueness, that they would not destroy this colour in their male offspring, their female offspring still inheriting the slaty tint. If so, the desired end of making a breed with the two sexes permanently different in colour might be gained.

The extreme importance, or rather necessity in the above case of the desired character, namely, pale-blueness, being present though in a latent state in the female, so that the male offspring should not be deteriorated, will be best appreciated as follows: the male of Soemmerring’s pheasant has a tail thirty-seven inches in length, whilst that of the female is only eight inches; the tail of the male common pheasant is about twenty inches, and that of the female twelve inches long. Now if the female Soemmerring pheasant with her SHORT tail were crossed with the male common pheasant, there can be no doubt that the male hybrid offspring would have a much LONGER tail than that of the pure offspring of the common pheasant. On the other hand, if the female common pheasant, with a tail much longer than that of the female Soemmerring pheasant, were crossed with the male of the latter, the male hybrid offspring would have a much SHORTER tail than that of the pure offspring of Soemmerring’s pheasant. (3. Temminck says that the tail of the female Phasianus Soemmerringii is only six inches long, ‘Planches coloriees,’ vol. v. 1838, pp. 487 and 488: the measurements above given were made for me by Mr. Sclater. For the common pheasant, see Macgillivray, ‘History of British Birds,’ vol. i. pp. 118-121.)

Our fancier, in order to make his new breed with the males of a pale-blue tint, and the females unchanged, would have to continue selecting the males during many generations; and each stage of paleness would have to be fixed in the males, and rendered latent in the females. The task would be an extremely difficult one, and has never been tried, but might possibly be successfully carried out. The chief obstacle would be the early and complete loss of the pale-blue tint, from the necessity of reiterated crosses with the slaty female, the latter not having at first any LATENT tendency to produce pale-blue offspring.

On the other hand, if one or two males were to vary ever so slightly in paleness, and the variations were from the first limited in their transmission to the male sex, the task of making a new breed of the desired kind would be easy, for such males would simply have to be selected and matched with ordinary females. An analogous case has actually occurred, for there are breeds of the pigeon in Belgium (4. Dr. Chapuis, ‘Le Pigeon Voyageur Belge,’ 1865, p. 87.) in which the males alone are marked with black striae. So again Mr. Tegetmeier has recently shewn (5. The ‘Field,’ Sept. 1872.) that dragons not rarely produce silver-coloured birds, which are almost always hens; and he himself has bred ten such females. It is on the other hand a very unusual event when a silver male is produced; so that nothing would be easier, if desired, than to make a breed of dragons with blue males and silver females. This tendency is indeed so strong that when Mr. Tegetmeier at last got a silver male and matched him with one of the silver females, he expected to get a breed with both sexes thus coloured; he was however disappointed, for the young male reverted to the blue colour of his grandfather, the young female alone being silver. No doubt with patience this tendency to reversion in the males, reared from an occasional silver male matched with a silver hen, might be eliminated, and then both sexes would be coloured alike; and this very process has been followed with success by Mr. Esquilant in the case of silver turbits.

With fowls, variations of colour, limited in their transmission to the male sex, habitually occur. When this form of inheritance prevails, it might well happen that some of the successive variations would be transferred to the female, who would then slightly resemble the male, as actually occurs in some breeds. Or again, the greater number, but not all, of the successive steps might be transferred to both sexes, and the female would then closely resemble the male. There can hardly be a doubt that this is the cause of the male pouter pigeon having a somewhat larger crop, and of the male carrier pigeon having somewhat larger wattles, than their respective females; for fanciers have not selected one sex more than the other, and have had no wish that these characters should be more strongly displayed in the male than in the female, yet this is the case with both breeds.

The same process would have to be followed, and the same difficulties encountered, if it were desired to make a breed with the females alone of some new colour.

Lastly, our fancier might wish to make a breed with the two sexes differing from each other, and both from the parent species. Here the difficulty would be extreme, unless the successive variations were from the first sexually limited on both sides, and then there would be no difficulty. We see this with the fowl; thus the two sexes of the pencilled Hamburghs differ greatly from each other, and from the two sexes of the aboriginal Gallus bankiva; and both are now kept constant to their standard of excellence by continued selection, which would be impossible unless the distinctive characters of both were limited in their transmission.

The Spanish fowl offers a more curious case; the male has an immense comb, but some of the successive variations, by the accumulation of which it was acquired, appear to have been transferred to the female; for she has a comb many times larger than that of the females of the parent species. But the comb of the female differs in one respect from that of the male, for it is apt to lop over; and within a recent period it has been ordered by the fancy that this should always be the case, and success has quickly followed the order. Now the lopping of the comb must be sexually limited in its transmission, otherwise it would prevent the comb of the male from being perfectly upright, which would be abhorrent to every fancier. On the other hand, the uprightness of the comb in the male must likewise be a sexually-limited character, otherwise it would prevent the comb of the female from lopping over.

From the foregoing illustrations, we see that even with almost unlimited time at command, it would be an extremely difficult and complex, perhaps an impossible process, to change one form of transmission into the other through selection. Therefore, without distinct evidence in each case, I am unwilling to admit that this has been effected in natural species. On the other hand, by means of successive variations, which were from the first

sexually limited in their transmission, there would not be the least difficulty in rendering a male bird widely different in colour or in any other character from the female; the latter being left unaltered, or slightly altered, or specially modified for the sake of protection.

As bright colours are of service to the males in their rivalry with other males, such colours would be selected whether or not they were transmitted exclusively to the same sex. Consequently the females might be expected often to partake of the brightness of the males to a greater or less degree; and this occurs with a host of species. If all the successive variations were transmitted equally to both sexes, the females would be indistinguishable from the males; and this likewise occurs with many birds. If, however, dull colours were of high importance for the safety of the female during incubation, as with many ground birds, the females which varied in brightness, or which received through inheritance from the males any marked accession of brightness, would sooner or later be destroyed. But the tendency in the males to continue for an indefinite period transmitting to their female offspring their own brightness, would have to be eliminated by a change in the form of inheritance; and this, as shewn by our previous illustration, would be extremely difficult. The more probable result of the long-continued destruction of the more brightly-coloured females, supposing the equal form of transmission to prevail, would be the lessening or annihilation of the bright colours of the males, owing to their continual crossing with the duller females. It would be tedious to follow out all the other possible results; but I may remind the reader that if sexually-limited variations in brightness occurred in the females, even if they were not in the least injurious to them and consequently were not eliminated, yet they would not be favoured or selected, for the male usually accepts any female, and does not select the more attractive individuals; consequently these variations would be liable to be lost, and would have little influence on the character of the race; and this will aid in accounting for the females being commonly duller-coloured than the males.

In the eighth chapter instances were given, to which many might here be added, of variations occurring at various ages, and inherited at the corresponding age. It was also shewn that variations which occur late in life are commonly transmitted to the same sex in which they first appear; whilst variations occurring early in life are apt to be transmitted to both sexes; not that all the cases of sexually-limited transmission can thus be accounted for. It was further shewn that if a male bird varied by becoming brighter whilst young, such variations would be of no service until the age for reproduction had arrived, and there was competition between rival males. But in the case of birds living on the ground and commonly in need of the protection of dull colours, bright tints would be far more dangerous to the young and inexperienced than to the adult males. Consequently the males which varied in brightness whilst young would suffer much destruction and be eliminated through natural selection; on the other hand, the males which varied in this manner when nearly mature, notwithstanding that they were exposed to some additional danger, might survive, and from being favoured through sexual selection, would procreate their kind. As a relation often exists between the period of variation and the form of transmission, if the bright-coloured young males were destroyed and the mature ones were successful in their courtship, the males alone would acquire brilliant colours and would transmit them exclusively to their male offspring. But I by no means wish to maintain that the influence of age on the form of transmission, is the sole cause of the great difference in brilliancy between the sexes of many birds.

When the sexes of birds differ in colour, it is interesting to determine whether the males alone have been modified by sexual selection, the females having been left unchanged, or only partially and indirectly thus changed; or whether the females have been specially modified through natural selection for the sake of protection. I will therefore discuss this question at some length, even more fully than its intrinsic importance deserves; for various curious collateral points may thus be conveniently considered.

Before we enter on the subject of colour, more especially in reference to Mr. Wallace’s conclusions, it may be useful to discuss some other sexual differences under a similar point of view. A breed of fowls formerly existed in Germany (6. Bechstein, ‘Naturgeschichte Deutschlands,’ 1793, B. iii. 339.) in which the hens were furnished with spurs; they were good layers, but they so greatly disturbed their nests with their spurs that they could not be allowed to sit on their own eggs. Hence at one time it appeared to me probable that with the females of the wild Gallinaceae the development of spurs had been checked through natural selection, from the injury thus caused to their nests. This seemed all the more probable, as wing-spurs, which would not be injurious during incubation, are often as well-developed in the female as in the male; though in not a few cases they are rather larger in the male. When the male is furnished with leg-spurs the female almost always exhibits rudiments of them,—the rudiment sometimes consisting of a mere scale, as in Gallus. Hence it might be argued that the females had aboriginally been furnished with well-developed spurs, but that these had subsequently been lost through disuse or natural selection. But if this view be admitted, it would have to be extended to innumerable other cases; and it implies that the female progenitors of the existing spur-bearing species were once encumbered with an injurious appendage.

In some few genera and species, as in Galloperdix, Acomus, and the Javan peacock (Pavo muticus), the females, as well as the males, possess well-developed leg-spurs. Are we to infer from this fact that they construct a different sort of nest from that made by their nearest allies, and not liable to be injured by their spurs; so that the spurs have not been removed? Or are we to suppose that the females of these several species especially require spurs for their defence? It is a more probable conclusion that both the presence and absence of spurs in the females result from different laws of inheritance having prevailed, independently of natural selection. With the many females in which spurs appear as rudiments, we may conclude that some few of the successive variations, through which they were developed in the males, occurred very early in life, and were consequently transferred to the females. In the other and much rarer cases, in which the females possess fully developed spurs, we may conclude that all the successive variations were transferred to them; and that they gradually acquired and inherited the habit of not disturbing their nests.

The vocal organs and the feathers variously modified for producing sound, as well as the proper instincts for using them, often differ in the two sexes, but are sometimes the same in both. Can such differences be accounted for by the males having acquired these organs and instincts, whilst the females have been saved from inheriting them, on account of the danger to which they would have been exposed by attracting the attention of birds or beasts of prey? This does not seem to me probable, when we think of the multitude of birds which with impunity gladden the country with their voices during the spring. (7. Daines Barrington, however, thought it probable (‘Philosophical Transactions,’ 1773, p. 164) that few female birds sing, because the talent would have been dangerous to them during incubation. He adds, that a similar view may possibly account for the inferiority of the female to the male in plumage.) It is a safer conclusion that, as vocal and instrumental organs are of special service only to the males during their courtship, these organs were developed through sexual selection and their constant use in that sex alone—the successive variations and the effects of use having been from the first more or less limited in transmission to the male offspring.

Many analogous cases could be adduced; those for instance of the plumes on the head being generally longer in the male than in the female, sometimes of equal length in both sexes, and occasionally absent in the female,—these several cases occurring in the same group of birds. It would be difficult to account for such a difference between the sexes by the female having been benefited by possessing a slightly shorter crest than the male, and its consequent diminution or complete suppression through natural selection. But I will take a more favourable case, namely the length of the tail. The long train of the peacock would have been not only inconvenient but dangerous to the peahen during the period of incubation and whilst accompanying her young. Hence there is not the least a priori improbability in the development of her tail having been checked through natural selection. But the females of various pheasants, which apparently are exposed on their open nests to as much danger as the peahen, have tails of considerable length. The females as well as the males of the Menura superba have long tails, and they build a domed nest, which is a great anomaly in so large a bird. Naturalists have wondered how the female Menura could manage her tail during incubation; but it is now known (8. Mr. Ramsay, in ‘Proc. Zoolog. Soc.’ 1868, p. 50.) that she “enters the nest head first, and then turns round with her tail sometimes over her back, but more often bent round by her side. Thus in time the tail becomes quite askew, and is a tolerable guide to the length of time the bird has been sitting.” Both sexes of an Australian kingfisher (Tanysiptera sylvia) have the middle tail-feathers greatly lengthened, and the female makes her nest in a hole; and as I am informed by Mr. R.B. Sharpe these feathers become much crumpled during incubation.

In these two latter cases the great length of the tail-feathers must be in some degree inconvenient to the female; and as in both species the tail-feathers of the female are somewhat shorter than those of the male, it might be argued that their full development had been prevented through natural selection. But if the development of the tail of the peahen had been checked only when it became inconveniently or dangerously great, she would have retained a much longer tail than she actually possesses; for her tail is not nearly so long, relatively to the size of her body, as that of many female pheasants, nor longer than that of the female turkey. It must also be borne in mind that, in accordance with this view, as soon as the tail of the peahen became dangerously long, and its development was consequently checked, she would have continually reacted on her male progeny, and thus have prevented the peacock from acquiring his present magnificent train. We may therefore infer that the length of the tail in the peacock and its shortness in the peahen are the result of the requisite variations in the male having been from the first transmitted to the male offspring alone.

We are led to a nearly similar conclusion with respect to the length of the tail in the various species of pheasants. In the Eared pheasant (Crossoptilon auritum) the tail is of equal length in both sexes, namely sixteen or seventeen inches; in the common pheasant it is about twenty inches long in the male and twelve in the female; in Soemmerring’s pheasant, thirty-seven inches in the male and only eight in the female; and lastly in Reeve’s pheasant it is sometimes actually seventy-two inches long in the male and sixteen in the female. Thus in the several species, the tail of the female differs much in length, irrespectively of that of the male; and this can be accounted for, as it seems to me, with much more probability, by the laws of inheritance,—that is by the successive variations having been from the first more or less closely limited in their transmission to the male sex than by the agency of natural selection, resulting from the length of tail being more or less injurious to the females of these several allied species.

We may now consider Mr. Wallace’s arguments in regard to the sexual coloration of birds. He believes that the bright tints originally acquired through sexual selection by the males would in all, or almost all cases, have been transmitted to the females, unless the transference had been checked through natural selection. I may here remind the reader that various facts opposed to this view have already been given under reptiles, amphibians, fishes and lepidoptera. Mr. Wallace rests his belief chiefly, but not exclusively, as we shall see in the next chapter, on the following statement (9. ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 78.), that when both sexes are coloured in a very conspicuous manner, the nest is of such a nature as to conceal the sitting bird; but when there is a marked contrast of colour between the sexes, the male being gay and the female dull-coloured, the nest is open and exposes the sitting bird to view. This coincidence, as far as it goes, certainly seems to favour the belief that the females which sit on open nests have been specially modified for the sake of protection; but we shall presently see that there is another and more probable explanation, namely, that conspicuous females have acquired the instinct of building domed nests oftener than dull-coloured birds. Mr. Wallace admits that there are, as might have been expected, some exceptions to his two rules, but it is a question whether the exceptions are not so numerous as seriously to invalidate them.

There is in the first place much truth in the Duke of Argyll’s remark (10. ‘Journal of Travel,’ edited by A. Murray, vol. i. 1868, p. 281.) that a large domed nest is more conspicuous to an enemy, especially to all tree-haunting carnivorous animals, than a smaller open nest. Nor must we forget that with many birds which build open nests, the male sits on the eggs and aids the female in feeding the young: this is the case, for instance, with Pyranga aestiva (11. Audubon, ‘Ornithological Biography,’ vol. i. p. 233.), one of the most splendid birds in the United States, the male being vermilion, and the female light brownish-green. Now if brilliant colours had been extremely dangerous to birds whilst sitting on their open nests, the males in these cases would have suffered greatly. It might, however, be of such paramount importance to the male to be brilliantly coloured, in order to beat his rivals, that this may have more than compensated some additional danger.

Mr. Wallace admits that with the King-crows (Dicrurus), Orioles, and Pittidae, the females are conspicuously coloured, yet build open nests; but he urges that the birds of the first group are highly pugnacious and could defend themselves; that those of the second group take extreme care in concealing their open nests, but this does not invariably hold good (12. Jerdon, ‘Birds of India,’ vol. ii. p. 108. Gould’s ‘Handbook of the Birds of Australia,’ vol. i. p. 463.); and that with the birds of the third group the females are brightly coloured chiefly on the under surface. Besides these cases, pigeons which are sometimes brightly, and almost always conspicuously coloured, and which are notoriously liable to the attacks of birds of prey, offer a serious exception to the rule, for they almost always build open and exposed nests. In another large family, that of the humming-birds, all the species build open nests, yet with some of the most gorgeous species the sexes are alike; and in the majority, the females, though less brilliant than the males, are brightly coloured. Nor can it be maintained that all female humming-birds, which are brightly coloured, escape detection by their tints being green, for some display on their upper surfaces red, blue, and other colours. (13. For instance, the female Eupetomena macroura has the head and tail dark blue with reddish loins; the female Lampornis porphyrurus is blackish-green on the upper surface, with the lores and sides of the throat crimson; the female Eulampis jugularis has the top of the head and back green, but the loins and the tail are crimson. Many other instances of highly conspicuous females could be given. See Mr. Gould’s magnificent work on this family.)

In regard to birds which build in holes or construct domed nests, other advantages, as Mr. Wallace remarks, besides concealment are gained, such as shelter from the rain, greater warmth, and in hot countries protection from the sun (14. Mr. Salvin noticed in Guatemala (‘Ibis,’ 1864, p. 375) that humming-birds were much more unwilling to leave their nests during very hot weather, when the sun was shining brightly, as if their eggs would be thus injured, than during cool, cloudy, or rainy weather.); so that it is no valid objection to his view that many birds having both sexes obscurely coloured build concealed nests. (15. I may specify, as instances of dull-coloured birds building concealed nests, the species belonging to eight Australian genera described in Gould’s ‘Handbook of the Birds of Australia,’ vol. i. pp. 340, 362, 365, 383, 387, 389, 391, 414.) The female Horn-bill (Buceros), for instance, of India and Africa is protected during incubation with extraordinary care, for she plasters up with her own excrement the orifice of the hole in which she sits on her eggs, leaving only a small orifice through which the male feeds her; she is thus kept a close prisoner during the whole period of incubation (16. Mr. C. Horne, ‘Proc. Zoolog. Soc.’ 1869. p. 243.); yet female horn-bills are not more conspicuously coloured than many other birds of equal size which build open nests. It is a more serious objection to Mr. Wallace’s view, as is admitted by him, that in some few groups the males are brilliantly coloured and the females obscure, and yet the latter hatch their eggs in domed nests. This is the case with the Grallinae of Australia, the Superb Warblers (Maluridae) of the same country, the Sun-birds (Nectariniae), and with several of the Australian Honey-suckers or Meliphagidae. (17. On the nidification and colours of these latter species, see Gould’s ‘Handbook to the Birds of Australia,’ vol. i. pp. 504, 527.)

If we look to the birds of England we shall see that there is no close and general relation between the colours of the female and the nature of the nest which is constructed. About forty of our British birds (excluding those of large size which could defend themselves) build in holes in banks, rocks, or trees, or construct domed nests. If we take the colours of the female goldfinch, bullfinch, or blackbird, as a standard of the degree of conspicuousness, which is not highly dangerous to the sitting female, then out of the above forty birds the females of only twelve can be considered as conspicuous to a dangerous degree, the remaining twenty-eight being inconspicuous. (18. I have consulted, on this subject, Macgillivray’s ‘British Birds,’ and though doubts may be entertained in some cases in regard to the degree of concealment of the nest, and to the degree of conspicuousness of the female, yet the following birds, which all lay their eggs in holes or in domed nests, can hardly be considered, by the above standard, as conspicuous: Passer, 2 species; Sturnus, of which the female is considerably less brilliant than the male; Cinclus; Motallica boarula (?); Erithacus (?); Fruticola, 2 sp.; Saxicola; Ruticilla, 2 sp.; Sylvia, 3 sp.; Parus, 3 sp.; Mecistura; Anorthura; Certhia; Sitta; Yunx; Muscicapa, 2 sp.; Hirundo, 3 sp.; and Cypselus. The females of the following 12 birds may be considered as conspicuous according to the same standard, viz., Pastor, Motacilla alba, Parus major and P. caeruleus, Upupa, Picus, 4 sp., Coracias, Alcedo, and Merops.) Nor is there any close relation within the same genus between a well-pronounced difference in colour between the sexes, and the nature of the nest constructed. Thus the male house sparrow (Passer domesticus) differs much from the female, the male tree-sparrow (P. montanus) hardly at all, and yet both build well-concealed nests. The two sexes of the common fly-catcher (Muscicapa grisola) can hardly be distinguished, whilst the sexes of the pied fly-catcher (M. luctuosa) differ considerably, and both species build in holes or conceal their nests. The female blackbird (Turdus merula) differs much, the female ring-ouzel (T. torquatus) differs less, and the female common thrush (T. musicus) hardly at all from their respective males; yet all build open nests. On the other hand, the not very distantly-allied water-ouzel (Cinclus aquaticus) builds a domed nest, and the sexes differ about as much as in the ring-ouzel. The black and red grouse (Tetrao tetrix and T. scoticus) build open nests in equally well-concealed spots, but in the one species the sexes differ greatly, and in the other very little.

Notwithstanding the foregoing objections, I cannot doubt, after reading Mr. Wallace’s excellent essay, that looking to the birds of the world, a large majority of the species in which the females are conspicuously coloured (and in this case the males with rare exceptions are equally conspicuous), build concealed nests for the sake of protection. Mr. Wallace enumerates (19. ‘Journal of Travel,’ edited by A. Murray, vol. i. p. 78.) a long series of groups in which this rule holds good; but it will suffice here to give, as instances, the more familiar groups of kingfishers, toucans, trogons, puff-birds (Capitonidae), plantain-eaters (Musophagae, woodpeckers, and parrots. Mr. Wallace believes that in these groups, as the males gradually acquired through sexual selection their brilliant colours, these were transferred to the females and were not eliminated by natural selection, owing to the protection which they already enjoyed from their manner of nidification. According to this view, their present manner of nesting was acquired before their present colours. But it seems to me much more probable that in most cases, as the females were gradually rendered more and more brilliant from partaking of the colours of the male, they were gradually led to change their instincts (supposing that they originally built open nests), and to seek protection by building domed or concealed nests. No one who studies, for instance, Audubon’s account of the differences in the nests of the same species in the Northern and Southern United States (20. See many statements in the ‘Ornithological Biography.’ See also some curious observations on the nests of Italian birds by Eugenio Bettoni, in the ‘Atti della Società Italiana,’ vol. xi. 1869, p. 487.), will feel any great difficulty in admitting that birds, either by a change (in the strict sense of the word) of their habits, or through the natural selection of so-called spontaneous variations of instinct, might readily be led to modify their manner of nesting.

This way of viewing the relation, as far as it holds good, between the bright colours of female birds and their manner of nesting, receives some support from certain cases occurring in the Sahara Desert. Here, as in most other deserts, various birds, and many other animals, have had their colours adapted in a wonderful manner to the tints of the surrounding surface. Nevertheless there are, as I am informed by the Rev. Mr. Tristram, some curious exceptions to the rule; thus the male of the Monticola cyanea is conspicuous from his bright blue colour, and the female almost equally conspicuous from her mottled brown and white plumage; both sexes of two species of Dromolaea are of a lustrous black; so that these three species are far from receiving protection from their colours, yet they are able to survive, for they have acquired the habit of taking refuge from danger in holes or crevices in the rocks.

With respect to the above groups in which the females are conspicuously coloured and build concealed nests, it is not necessary to suppose that each separate species had its nidifying instinct specially modified; but only that the early progenitors of each group were gradually led to build domed or concealed nests, and afterwards transmitted this instinct, together with their bright colours, to their modified descendants. As far as it can be trusted, the conclusion is interesting, that sexual selection together with equal or nearly equal inheritance by both sexes, have indirectly determined the manner of nidification of whole groups of birds.

According to Mr. Wallace, even in the groups in which the females, from being protected in domed nests during incubation, have not had their bright colours eliminated through natural selection, the males often differ in a slight, and occasionally in a considerable degree from the females. This is a significant fact, for such differences in colour must be accounted for by some of the variations in the males having been from the first limited in transmission to the same sex; as it can hardly be maintained that these differences, especially when very slight, serve as a protection to the female. Thus all the species in the splendid group of the Trogons build in holes; and Mr. Gould gives figures (21. See his Monograph of the Trogonidae, 1st edition.) of both sexes of twenty-five species, in all of which, with one partial exception, the sexes differ sometimes slightly, sometimes conspicuously, in colour,—the males being always finer than the females, though the latter are likewise beautiful. All the species of kingfishers build in holes, and with most of the species the sexes are equally brilliant, and thus far Mr. Wallace’s rule holds good; but in some of the Australian species the colours of the females are rather less vivid than those of the male; and in one splendidly-coloured species, the sexes differ so much that they were at first thought to be specifically distinct. (22. Namely, Cyanalcyon, Gould’s ‘Handbook to the Birds of Australia,’ vol. i. p. 133; see, also, pp. 130, 136.) Mr. R.B. Sharpe, who has especially studied this group, has shewn me some American species (Ceryle) in which the breast of the male is belted with black. Again, in Carcineutes, the difference between the sexes is conspicuous: in the male the upper surface is dull-blue banded with black, the lower surface being partly fawn-coloured, and there is much red about the head; in the female the upper surface is reddish-brown banded with black, and the lower surface white with black markings. It is an interesting fact, as shewing how the same peculiar style of sexual colouring often characterises allied forms, that in three species of Dacelo the male differs from the female only in the tail being dull-blue banded with black, whilst that of the female is brown with blackish bars; so that here the tail differs in colour in the two sexes in exactly the same manner as the whole upper surface in the two sexes of Carcineutes.

With parrots, which likewise build in holes, we find analogous cases: in most of the species, both sexes are brilliantly coloured and indistinguishable, but in not a few species the males are coloured rather more vividly than the females, or even very differently from them. Thus, besides other strongly-marked differences, the whole under surface of the male King Lory (Aprosmictus scapulatus) is scarlet, whilst the throat and chest of the female is green tinged with red: in the Euphema splendida there is a similar difference, the face and wing coverts moreover of the female being of a paler blue than in the male. (23. Every gradation of difference between the sexes may be followed in the parrots of Australia. See Gould’s ‘Handbook,’ etc., vol. ii. pp. 14-102.) In the family of the tits (Parinae), which build concealed nests, the female of our common blue tomtit (Parus caeruleus), is “much less brightly coloured” than the male: and in the magnificent Sultan yellow tit of India the difference is greater. (24. Macgillivray’s ‘British Birds,’ vol. ii. p. 433. Jerdon, ‘Birds of India,’ vol. ii. p. 282.)

Again, in the great group of the woodpeckers (25. All the following facts are taken from M. Malherbe’s magnificent ‘Monographie des Picidees,’ 1861.), the sexes are generally nearly alike, but in the Megapicus validus all those parts of the head, neck, and breast, which are crimson in the male are pale brown in the female. As in several woodpeckers the head of the male is bright crimson, whilst that of the female is plain, it occurred to me that this colour might possibly make the female dangerously conspicuous, whenever she put her head out of the hole containing her nest, and consequently that this colour, in accordance with Mr. Wallace’s belief, had been eliminated. This view is strengthened by what Malherbe states with respect to Indopicus carlotta; namely, that the young females, like the young males, have some crimson about their heads, but that this colour disappears in the adult female, whilst it is intensified in the adult male. Nevertheless the following considerations render this view extremely doubtful: the male takes a fair share in incubation (26. Audubon’s ‘Ornithological Biography,’ vol. ii. p. 75; see also the ‘Ibis,’ vol. i. p. 268.), and would be thus almost equally exposed to danger; both sexes of many species have their heads of an equally bright crimson; in other species the difference between the sexes in the amount of scarlet is so slight that it can hardly make any appreciable difference in the danger incurred; and lastly, the colouring of the head in the two sexes often differs slightly in other ways.

The cases, as yet given, of slight and graduated differences in colour between the males and females in the groups, in which as a general rule the sexes resemble each other, all relate to species which build domed or concealed nests. But similar gradations may likewise be observed in groups in which the sexes as a general rule resemble each other, but which build open nests.

As I have before instanced the Australian parrots, so I may here instance, without giving any details, the Australian pigeons. (27. Gould’s ‘Handbook to the Birds of Australia,’ vol. ii. pp. 109-149.) It deserves especial notice that in all these cases the slight differences in plumage between the sexes are of the same general nature as the occasionally greater differences. A good illustration of this fact has already been afforded by those kingfishers in which either the tail alone or the whole upper surface of the plumage differs in the same manner in the two sexes. Similar cases may be observed with parrots and pigeons. The differences in colour between the sexes of the same species are, also, of the same general nature as the differences in colour between the distinct species of the same group. For when in a group in which the sexes are usually alike, the male differs considerably from the female, he is not coloured in a quite new style. Hence we may infer that within the same group the special colours of both sexes when they are alike, and the colours of the male, when he differs slightly or even considerably from the female, have been in most cases determined by the same general cause; this being sexual selection.

It is not probable, as has already been remarked, that differences in colour between the sexes, when very slight, can be of service to the female as a protection. Assuming, however, that they are of service, they might be thought to be cases of transition; but we have no reason to believe that many species at any one time are undergoing change. Therefore we can hardly admit that the numerous females which differ very slightly in colour from their males are now all commencing to become obscure for the sake of protection. Even if we consider somewhat more marked sexual differences, is it probable, for instance, that the head of the female chaffinch,—the crimson on the breast of the female bullfinch,—the green of the female greenfinch,—the crest of the female golden-crested wren, have all been rendered less bright by the slow process of selection for the sake of protection? I cannot think so; and still less with the slight differences between the sexes of those birds which build concealed nests. On the other hand, the differences in colour between the sexes, whether great or small, may to a large extent be explained on the principle of the successive variations, acquired by the males through sexual selection, having been from the first more or less limited in their transmission to the females. That the degree of limitation should differ in different species of the same group will not surprise any one who has studied the laws of inheritance, for they are so complex that they appear to us in our ignorance to be capricious in their action. (28. See remarks to this effect in ‘Variation of Animals and Plants under Domestication,’ vol. ii. chap. xii.)

As far as I can discover there are few large groups of birds in which all the species have both sexes alike and brilliantly coloured, but I hear from Mr. Sclater, that this appears to be the case with the Musophagae or plantain-eaters. Nor do I believe that any large group exists in which the sexes of all the species are widely dissimilar in colour: Mr. Wallace informs me that the chatterers of S. America (Cotingidae) offer one of the best instances; but with some of the species, in which the male has a splendid red breast, the female exhibits some red on her breast; and the females of other species shew traces of the green and other colours of the males. Nevertheless we have a near approach to close sexual similarity or dissimilarity throughout several groups: and this, from what has just been said of the fluctuating nature of inheritance, is a somewhat surprising circumstance. But that the same laws should largely prevail with allied animals is not surprising. The domestic fowl has produced a great number of breeds and sub-breeds, and in these the sexes generally differ in plumage; so that it has been noticed as an unusual circumstance when in certain sub-breeds they resemble each other. On the other hand, the domestic pigeon has likewise produced a vast number of distinct breeds and sub-breeds, and in these, with rare exceptions, the two sexes are identically alike.

Therefore if other species of Gallus and Columba were domesticated and varied, it would not be rash to predict that similar rules of sexual similarity and dissimilarity, depending on the form of transmission, would hold good in both cases. In like manner the same form of transmission has generally prevailed under nature throughout the same groups, although marked exceptions to this rule occur. Thus within the same family or even genus, the sexes may be identically alike, or very different in colour. Instances have already been given in the same genus, as with sparrows, fly-catchers, thrushes and grouse. In the family of pheasants the sexes of almost all the species are wonderfully dissimilar, but are quite alike in the eared pheasant or Crossoptilon auritum. In two species of Chloephaga, a genus of geese, the male cannot be distinguished from the females, except by size; whilst in two others, the sexes are so unlike that they might easily be mistaken for distinct species. (29. The ‘Ibis,’ vol. vi. 1864, p. 122.)

The laws of inheritance can alone account for the following cases, in which the female acquires, late in life, certain characters proper to the male, and ultimately comes to resemble him more or less completely. Here protection can hardly have come into play. Mr. Blyth informs me that the females of Oriolus melanocephalus and of some allied species, when sufficiently mature to breed, differ considerably in plumage from the adult males; but after the second or third moults they differ only in their beaks having a slight greenish tinge. In the dwarf bitterns (Ardetta), according to the same authority, “the male acquires his final livery at the first moult, the female not before the third or fourth moult; in the meanwhile she presents an intermediate garb, which is ultimately exchanged for the same livery as that of the male.” So again the female Falco peregrinus acquires her blue plumage more slowly than the male. Mr. Swinhoe states that with one of the Drongo shrikes (Dicrurus macrocercus) the male, whilst almost a nestling, moults his soft brown plumage and becomes of a uniform glossy greenish-black; but the female retains for a long time the white striae and spots on the axillary feathers; and does not completely assume the uniform black colour of the male for three years. The same excellent observer remarks that in the spring of the second year the female spoon-bill (Platalea) of China resembles the male of the first year, and that apparently it is not until the third spring that she acquires the same adult plumage as that possessed by the male at a much earlier age. The female Bombycilla carolinensis differs very little from the male, but the appendages, which like beads of red sealing-wax ornament the wing-feathers (30. When the male courts the female, these ornaments are vibrated, and “are shewn off to great advantage,” on the outstretched wings: A. Leith Adams, ‘Field and Forest Rambles,’ 1873, p. 153.), are not developed in her so early in life as in the male. In the male of an Indian parrakeet (Palaeornis javanicus) the upper mandible is coral-red from his earliest youth, but in the female, as Mr. Blyth has observed with caged and wild birds, it is at first black and does not become red until the bird is at least a year old, at which age the sexes resemble each other in all respects. Both sexes of the wild turkey are ultimately furnished with a tuft of bristles on the breast, but in two-year-old birds the tuft is about four inches long in the male and hardly apparent in the female; when, however, the latter has reached her fourth year, it is from four to five inches in length. (31. On Ardetta, Translation of Cuvier’s ‘Regne Animal,’ by Mr. Blyth, footnote, p. 159. On the Peregrine Falcon, Mr. Blyth, in Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 304. On Dicrurus, ‘Ibis,’ 1863, p. 44. On the Platalea, ‘Ibis,’ vol. vi. 1864, p. 366. On the Bombycilla, Audubon’s ‘Ornitholog. Biography,’ vol. i. p. 229. On the Palaeornis, see, also, Jerdon, ‘Birds of India,’ vol. i. p. 263. On the wild turkey, Audubon, ibid. vol. i. p. 15; but I hear from Judge Caton that in Illinois the female very rarely acquires a tuft. Analogous cases with the females of Petrocossyphus are given by Mr. R. Sharpe, ‘Proceedings of the Zoological Society,’ 1872, p. 496.)

These cases must not be confounded with those where diseased or old females abnormally assume masculine characters, nor with those where fertile females, whilst young, acquire the characters of the male, through variation or some unknown cause. (32. Of these latter cases Mr. Blyth has recorded (Translation of Cuvier’s ‘Regne Animal,’ p. 158) various instances with Lanius, Ruticilla, Linaria, and Anas. Audubon has also recorded a similar case (‘Ornitholog. Biography,’ vol. v. p. 519) with Pyranga aestiva.) But all these cases have so much in common that they depend, according to the hypothesis of pangenesis, on gemmules derived from each part of the male being present, though latent, in the female; their development following on some slight change in the elective affinities of her constituent tissues.

A few words must be added on changes of plumage in relation to the season of the year. From reasons formerly assigned there can be little doubt that the elegant plumes, long pendant feathers, crests, etc., of egrets, herons, and many other birds, which are developed and retained only during the summer, serve for ornamental and nuptial purposes, though common to both sexes. The female is thus rendered more conspicuous during the period of incubation than during the winter; but such birds as herons and egrets would be able to defend themselves. As, however, plumes would probably be inconvenient and certainly of no use during the winter, it is possible that the habit of moulting twice in the year may have been gradually acquired through natural selection for the sake of casting off inconvenient ornaments during the winter. But this view cannot be extended to the many waders, whose summer and winter plumages differ very little in colour. With defenceless species, in which both sexes, or the males alone, become extremely conspicuous during the breeding-season,—or when the males acquire at this season such long wing or tail-feathers as to impede their flight, as with Cosmetornis and Vidua,—it certainly at first appears highly probable that the second moult has been gained for the special purpose of throwing off these ornaments. We must, however, remember that many birds, such as some of the Birds of Paradise, the Argus pheasant and peacock, do not cast their plumes during the winter; and it can hardly be maintained that the constitution of these birds, at least of the Gallinaceae, renders a double moult impossible, for the ptarmigan moults thrice in the year. (33. See Gould’s ‘Birds of Great Britain.’) Hence it must be considered as doubtful whether the many species which moult their ornamental plumes or lose their bright colours during the winter, have acquired this habit on account of the inconvenience or danger which they would otherwise have suffered.

I conclude, therefore, that the habit of moulting twice in the year was in most or all cases first acquired for some distinct purpose, perhaps for gaining a warmer winter covering; and that variations in the plumage occurring during the summer were accumulated through sexual selection, and transmitted to the offspring at the same season of the year; that such variations were inherited either by both sexes or by the males alone, according to the form of inheritance which prevailed. This appears more probable than that the species in all cases originally tended to retain their ornamental plumage during the winter, but were saved from this through natural selection, resulting from the inconvenience or danger thus caused.

I have endeavoured in this chapter to shew that the arguments are not trustworthy in favour of the view that weapons, bright colours, and various ornaments, are now confined to the males owing to the conversion, by natural selection, of the equal transmission of characters to both sexes, into transmission to the male sex alone. It is also doubtful whether the colours of many female birds are due to the preservation, for the sake of protection, of variations which were from the first limited in their transmission to the female sex. But it will be convenient to defer any further discussion on this subject until I treat, in the following chapter, of the differences in plumage between the young and old.

XVI. BIRDS—concluded

The immature plumage in relation to the character of the plumage in both sexes when adult—Six classes of cases—Sexual differences between the males of closely-allied or representative species—The female assuming the characters of the male—Plumage of the young in relation to the summer and winter plumage of the adults—On the increase of beauty in the birds of the world—Protective colouring—Conspicuously coloured birds—Novelty appreciated—Summary of the four chapters on Birds.

We must now consider the transmission of characters, as limited by age, in reference to sexual selection. The truth and importance of the principle of inheritance at corresponding ages need not here be discussed, as enough has already been said on the subject. Before giving the several rather complex rules or classes of cases, under which the differences in plumage between the young and the old, as far as known to me, may be included, it will be well to make a few preliminary remarks.

With animals of all kinds when the adults differ in colour from the young, and the colours of the latter are not, as far as we can see, of any special service, they may generally be attributed, like various embryological structures, to the retention of a former character. But this view can be maintained with confidence, only when the young of several species resemble each other closely, and likewise resemble other adult species belonging to the same group; for the latter are the living proofs that such a state of things was formerly possible. Young lions and pumas are marked with feeble stripes or rows of spots, and as many allied species both young and old are similarly marked, no believer in evolution will doubt that the progenitor of the lion and puma was a striped animal, and that the young have retained vestiges of the stripes, like the kittens of black cats, which are not in the least striped when grown up. Many species of deer, which when mature are not spotted, are whilst young covered with white spots, as are likewise some few species in the adult state. So again the young in the whole family of pigs (Suidae), and in certain rather distantly allied animals, such as the tapir, are marked with dark longitudinal stripes; but here we have a character apparently derived from an extinct progenitor, and now preserved by the young alone. In all such cases the old have had their colours changed in the course of time, whilst the young have remained but little altered, and this has been effected through the principle of inheritance at corresponding ages.

This same principle applies to many birds belonging to various groups, in which the young closely resemble each other, and differ much from their respective adult parents. The young of almost all the Gallinaceae, and of some distantly allied birds such as ostriches, are covered with longitudinally striped down; but this character points back to a state of things so remote that it hardly concerns us. Young cross-bills (Loxia) have at first straight beaks like those of other finches, and in their immature striated plumage they resemble the mature red-pole and female siskin, as well as the young of the goldfinch, greenfinch, and some other allied species. The young of many kinds of buntings (Emberiza) resemble one another, and likewise the adult state of the common bunting, E. miliaria. In almost the whole large group of thrushes the young have their breasts spotted—a character which is retained throughout life by many species, but is quite lost by others, as by the Turdus migratorius. So again with many thrushes, the feathers on the back are mottled before they are moulted for the first time, and this character is retained for life by certain eastern species. The young of many species of shrikes (Lanius), of some woodpeckers, and of an Indian pigeon (Chalcophaps indicus), are transversely striped on the under surface; and certain allied species or whole genera are similarly marked when adult. In some closely-allied and resplendent Indian cuckoos (Chrysococcyx), the mature species differ considerably from one another in colour, but the young cannot be distinguished. The young of an Indian goose (Sarkidiornis melanonotus) closely resemble in plumage an allied genus, Dendrocygna, when mature. (1. In regard to thrushes, shrikes, and woodpeckers, see Mr. Blyth, in Charlesworth’s ‘Mag. of Nat. Hist.’ vol. i. 1837, p. 304; also footnote to his translation of Cuvier’s ‘Regne Animal,’ p. 159. I give the case of Loxia on Mr. Blyth’s information. On thrushes, see also Audubon, ‘Ornith. Biog.’ vol. ii. p. 195. On Chrysococcyx and Chalcophaps, Blyth, as quoted in Jerdon’s ‘Birds of India,’ vol. iii. p. 485. On Sarkidiornis, Blyth, in ‘Ibis,’ 1867, p. 175.) Similar facts will hereafter be given in regard to certain herons. Young black-grouse (Tetrao tetrix) resemble the young as well as the old of certain other species, for instance the red-grouse or T. scoticus. Finally, as Mr. Blyth, who has attended closely to this subject, has well remarked, the natural affinities of many species are best exhibited in their immature plumage; and as the true affinities of all organic beings depend on their descent from a common progenitor, this remark strongly confirms the belief that the immature plumage approximately shews us the former or ancestral condition of the species.

Although many young birds, belonging to various families, thus give us a glimpse of the plumage of their remote progenitors, yet there are many other birds, both dull-coloured and bright-coloured, in which the young closely resemble their parents. In such cases the young of the different species cannot resemble each other more closely than do the parents; nor can they strikingly resemble allied forms when adult. They give us but little insight into the plumage of their progenitors, excepting in so far that, when the young and the old are coloured in the same general manner throughout a whole group of species, it is probable that their progenitors were similarly coloured.

We may now consider the classes of cases, under which the differences and resemblances between the plumage of the young and the old, in both sexes or in one sex alone, may be grouped. Rules of this kind were first enounced by Cuvier; but with the progress of knowledge they require some modification and amplification. This I have attempted to do, as far as the extreme complexity of the subject permits, from information derived from various sources; but a full essay on this subject by some competent ornithologist is much needed. In order to ascertain to what extent each rule prevails, I have tabulated the facts given in four great works, namely, by Macgillivray on the birds of Britain, Audubon on those of North America, Jerdon on those of India, and Gould on those of Australia. I may here premise, first, that the several cases or rules graduate into each other; and secondly, that when the young are said to resemble their parents, it is not meant that they are identically alike, for their colours are almost always less vivid, and the feathers are softer and often of a different shape.

RULES OR CLASSES OF CASES.

I. When the adult male is more beautiful or conspicuous than the adult female, the young of both sexes in their first plumage closely resemble the adult female, as with the common fowl and peacock; or, as occasionally occurs, they resemble her much more closely than they do the adult male.

II. When the adult female is more conspicuous than the adult male, as sometimes though rarely occurs, the young of both sexes in their first plumage resemble the adult male.

III. When the adult male resembles the adult female, the young of both sexes have a peculiar first plumage of their own, as with the robin.

IV. When the adult male resembles the adult female, the young of both sexes in their first plumage resemble the adults, as with the kingfisher, many parrots, crows, hedge-warblers.

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.

VI. In some few cases 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.

CLASS I. — In this class, the young of both sexes more or less closely resemble the adult female, whilst the adult male differs from the adult female, often in the most conspicuous manner. Innumerable instances in all Orders could be given; it will suffice to call to mind the common pheasant, duck, and house-sparrow. The cases under this class graduate into others. Thus the two sexes when adult may differ so slightly, and the young so slightly from the adults, that it is doubtful whether such cases ought to come under the present, or under the third or fourth classes. So again the young of the two sexes, instead of being quite alike, may differ in a slight degree from each other, as in our sixth class. These transitional cases, however, are few, or at least are not strongly pronounced, in comparison with those which come strictly under the present class.

The force of the present law is well shewn in those groups, in which, as a general rule, the two sexes and the young are all alike; for when in these groups the male does differ from the female, as with certain parrots, kingfishers, pigeons, etc., the young of both sexes resemble the adult female. (2. See, for instance, Mr. Gould’s account (‘Handbook to the Birds of Australia,’ vol. i. p. 133) of Cyanalcyon (one of the Kingfishers), in which, however, the young male, though resembling the adult female, is less brilliantly coloured. In some species of Dacelo the males have blue tails, and the females brown ones; and Mr. R.B. Sharpe informs me that the tail of the young male of D. gaudichaudi is at first brown. Mr. Gould has described (ibid. vol. ii. pp. 14, 20, 37) the sexes and the young of certain black Cockatoos and of the King Lory, with which the same rule prevails. Also Jerdon (‘Birds of India,’ vol. i. p. 260) on the Palaeornis rosa, in which the young are more like the female than the male. See Audubon (‘Ornithological Biography,’ vol. ii. p. 475) on the two sexes and the young of Columba passerina.) We see the same fact exhibited still more clearly in certain anomalous cases; thus the male of Heliothrix auriculata (one of the humming-birds) differs conspicuously from the female in having a splendid gorget and fine ear-tufts, but the female is remarkable from having a much longer tail than that of the male; now the young of both sexes resemble (with the exception of the breast being spotted with bronze) the adult female in all other respects, including the length of her tail, so that the tail of the male actually becomes shorter as he reaches maturity, which is a most unusual circumstance. (3. I owe this information to Mr. Gould, who shewed me the specimens; see also his ‘Introduction to the Trochilidae,’ 1861, p. 120.) Again, the plumage of the male goosander (Mergus merganser) is more conspicuously coloured than that of the female, with the scapular and secondary wing-feathers much longer; but differently from what occurs, as far as I know, in any other bird, the crest of the adult male, though broader than that of the female, is considerably shorter, being only a little above an inch in length; the crest of the female being two and a half inches long. Now the young of both sexes entirely resemble the adult female, so that their crests are actually of greater length, though narrower, than in the adult male. (4. Macgillivray, ‘Hist. Brit. Birds,’ vol. v. pp. 207-214.)

When the young and the females closely resemble each other and both differ from the males, the most obvious conclusion is that the males alone have been modified. Even in the anomalous cases of the Heliothrix and Mergus, it is probable that originally both adult sexes were furnished—the one species with a much elongated tail, and the other with a much elongated crest—these characters having since been partially lost by the adult males from some unexplained cause, and transmitted in their diminished state to their male offspring alone, when arrived at the corresponding age of maturity. The belief that in the present class the male alone has been modified, as far as the differences between the male and the female together with her young are concerned, is strongly supported by some remarkable facts recorded by Mr. Blyth (5. See his admirable paper in the ‘Journal of the Asiatic Soc. of Bengal,’ vol. xix. 1850, p. 223; see also Jerdon, ‘Birds of India,’ vol. i. introduction, p. xxix. In regard to Tanysiptera, Prof. Schlegel told Mr. Blyth that he could distinguish several distinct races, solely by comparing the adult males.), with respect to closely-allied species which represent each other in distinct countries. For with several of these representative species the adult males have undergone a certain amount of change and can be distinguished; the females and the young from the distinct countries being indistinguishable, and therefore absolutely unchanged. This is the case with certain Indian chats (Thamnobia), with certain honey-suckers (Nectarinia), shrikes (Tephrodornis), certain kingfishers (Tanysiptera), Kalij pheasants (Gallophasis), and tree-partridges (Arboricola).

In some analogous cases, namely with birds having a different summer and winter plumage, but with the two sexes nearly alike, certain closely-allied species can easily be distinguished in their summer or nuptial plumage, yet are indistinguishable in their winter as well as in their immature plumage. This is the case with some of the closely-allied Indian wagtails or Motacillae. Mr. Swinhoe (6. See also Mr. Swinhoe, in ‘Ibis,’ July 1863, p. 131; and a previous paper, with an extract from a note by Mr. Blyth, in ‘Ibis,’ January, 1861, p. 25.) informs me that three species of Ardeola, a genus of herons, which represent one another on separate continents, are “most strikingly different” when ornamented with their summer plumes, but are hardly, if at all, distinguishable during the winter. The young also of these three species in their immature plumage closely resemble the adults in their winter dress. This case is all the more interesting, because with two other species of Ardeola both sexes retain, during the winter and summer, nearly the same plumage as that possessed by the three first species during the winter and in their immature state; and this plumage, which is common to several distinct species at different ages and seasons, probably shews us how the progenitors of the genus were coloured. In all these cases, the nuptial plumage which we may assume was originally acquired by the adult males during the breeding-season, and transmitted to the adults of both sexes at the corresponding season, has been modified, whilst the winter and immature plumages have been left unchanged.

The question naturally arises, how is it that in these latter cases the winter plumage of both sexes, and in the former cases the plumage of the adult females, as well as the immature plumage of the young, have not been at all affected? The species which represent each other in distinct countries will almost always have been exposed to somewhat different conditions, but we can hardly attribute to this action the modification of the plumage in the males alone, seeing that the females and the young, though similarly exposed, have not been affected. Hardly any fact shews us more clearly how subordinate in importance is the direct action of the conditions of life, in comparison with the accumulation through selection of indefinite variations, than the surprising difference between the sexes of many birds; for both will have consumed the same food, and have been exposed to the same climate. Nevertheless we are not precluded from believing that in the course of time new conditions may produce some direct effect either on both sexes, or from their constitutional differences chiefly on one sex. We see only that this is subordinate in importance to the accumulated results of selection. Judging, however, from a wide-spread analogy, when a species migrates into a new country (and this must precede the formation of representative species), the changed conditions to which they will almost always have been exposed will cause them to undergo a certain amount of fluctuating variability. In this case sexual selection, which depends on an element liable to change—the taste or admiration of the female—will have had new shades of colour or other differences to act on and accumulate; and as sexual selection is always at work, it would (from what we know of the results on domestic animals of man’s unintentional selection), be surprising if animals inhabiting separate districts, which can never cross and thus blend their newly-acquired characters, were not, after a sufficient lapse of time, differently modified. These remarks likewise apply to the nuptial or summer plumage, whether confined to the males, or common to both sexes.

Although the females of the above closely-allied or representative species, together with their young, differ hardly at all from one another, so that the males alone can be distinguished, yet the females of most species within the same genus obviously differ from each other. The differences, however, are rarely as great as between the males. We see this clearly in the whole family of the Gallinaceae: the females, for instance, of the common and Japan pheasant, and especially of the gold and Amherst pheasant —of the silver pheasant and the wild fowl—resemble one another very closely in colour, whilst the males differ to an extraordinary degree. So it is with the females of most of the Cotingidae, Fringillidae, and many other families. There can indeed be no doubt that, as a general rule, the females have been less modified than the males. Some few birds, however, offer a singular and inexplicable exception; thus the females of Paradisea apoda and P. papuana differ from each other more than do their respective males (7. Wallace, ‘The Malay Archipelago,’ vol. ii. 1869, p. 394.); the female of the latter species having the under surface pure white, whilst the female P. apoda is deep brown beneath. So, again, as I hear from Professor Newton, the males of two species of Oxynotus (shrikes), which represent each other in the islands of Mauritius and Bourbon (8. These species are described with coloured figures, by M. F. Pollen, in ‘Ibis,’ 1866, p. 275.), differ but little in colour, whilst the females differ much. In the Bourbon species the female appears to have partially retained an immature condition of plumage, for at first sight she “might be taken for the young of the Mauritian species.” These differences may be compared with those inexplicable ones, which occur independently of man’s selection in certain sub-breeds of the game-fowl, in which the females are very different, whilst the males can hardly be distinguished. (9. ‘Variation of Animals,’ etc., vol. i. p. 251.)

As I account so largely by sexual selection for the differences between the males of allied species, how can the differences between the females be accounted for in all ordinary cases? We need not here consider the species which belong to distinct genera; for with these, adaptation to different habits of life, and other agencies, will have come into play. In regard to the differences between the females within the same genus, it appears to me almost certain, after looking through various large groups, that the chief agent has been the greater or less transference to the female of the characters acquired by the males through sexual selection. In the several British finches, the two sexes differ either very slightly or considerably; and if we compare the females of the greenfinch, chaffinch, goldfinch, bullfinch, crossbill, sparrow, etc., we shall see that they differ from one another chiefly in the points in which they partially resemble their respective males; and the colours of the males may safely be attributed to sexual selection. With many gallinaceous species the sexes differ to an extreme degree, as with the peacock, pheasant, and fowl, whilst with other species there has been a partial or even complete transference of character from the male to the female. The females of the several species of Polyplectron exhibit in a dim condition, and chiefly on the tail, the splendid ocelli of their males. The female partridge differs from the male only in the red mark on her breast being smaller; and the female wild turkey only in her colours being much duller. In the guinea-fowl the two sexes are indistinguishable. There is no improbability in the plain, though peculiarly spotted plumage of this latter bird having been acquired through sexual selection by the males, and then transmitted to both sexes; for it is not essentially different from the much more beautifully spotted plumage, characteristic of the males alone of the Tragopan pheasants.

It should be observed that, in some instances, the transference of characters from the male to the female has been effected apparently at a remote period, the male having subsequently undergone great changes, without transferring to the female any of his later-gained characters. For instance, the female and the young of the black-grouse (Tetrao tetrix) resemble pretty closely both sexes and the young of the red-grouse (T. scoticus); and we may consequently infer that the black-grouse is descended from some ancient species, of which both sexes were coloured in nearly the same manner as the red-grouse. As both sexes of this latter species are more distinctly barred during the breeding-season than at any other time, and as the male differs slightly from the female in his more strongly-pronounced red and brown tints (10. Macgillivray, ‘History of British Birds,’ vol. i. pp. 172-174.), we may conclude that his plumage has been influenced by sexual selection, at least to a certain extent. If so, we may further infer that nearly similar plumage of the female black-grouse was similarly produced at some former period. But since this period the male black-grouse has acquired his fine black plumage, with his forked and outwardly-curled tail-feathers; but of these characters there has hardly been any transference to the female, excepting that she shews in her tail a trace of the curved fork.

We may therefore conclude that the females of distinct though allied species have often had their plumage rendered more or less different by the transference in various degrees of characters acquired by the males through sexual selection, both during former and recent times. But it deserves especial attention that brilliant colours have been transferred much more rarely than other tints. For instance, the male of the red-throated blue-breast (Cyanecula suecica) has a rich blue breast, including a sub-triangular red mark; now marks of nearly the same shape have been transferred to the female, but the central space is fulvous instead of red, and is surrounded by mottled instead of blue feathers. The Gallinaceae offer many analogous cases; for none of the species, such as partridges, quails, guinea-fowls, etc., in which the colours of the plumage have been largely transferred from the male to the female, are brilliantly coloured. This is well exemplified with the pheasants, in which the male is generally so much more brilliant than the female; but with the Eared and Cheer pheasants (Crossoptilon auritum and Phasianus wallichii) the sexes closely resemble each other and their colours are dull. We may go so far as to believe that if any part of the plumage in the males of these two pheasants had been brilliantly coloured, it would not have been transferred to the females. These facts strongly support Mr. Wallace’s view that with birds which are exposed to much danger during incubation, the transference of bright colours from the male to the female has been checked through natural selection. We must not, however, forget that another explanation, before given, is possible; namely, that the males which varied and became bright, whilst they were young and inexperienced, would have been exposed to much danger, and would generally have been destroyed; the older and more cautious males, on the other hand, if they varied in a like manner, would not only have been able to survive, but would have been favoured in their rivalry with other males. Now variations occurring late in life tend to be transmitted exclusively to the same sex, so that in this case extremely bright tints would not have been transmitted to the females. On the other hand, ornaments of a less conspicuous kind, such as those possessed by the Eared and Cheer pheasants, would not have been dangerous, and if they appeared during early youth, would generally have been transmitted to both sexes.

In addition to the effects of the partial transference of characters from the males to the females, some of the differences between the females of closely allied species may be attributed to the direct or definite action of the conditions of life. (11. See, on this subject, chap. xxiii. in the ‘Variation of Animals and Plants under Domestication.’) With the males, any such action would generally have been masked by the brilliant colours gained through sexual selection; but not so with the females. Each of the endless diversities in plumage which we see in our domesticated birds is, of course, the result of some definite cause; and under natural and more uniform conditions, some one tint, assuming that it was in no way injurious, would almost certainly sooner or later prevail. The free intercrossing of the many individuals belonging to the same species would ultimately tend to make any change of colour, thus induced, uniform in character.

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).)

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. (33. I am indebted to Mr. Blyth for information in regard to this genus. The sparrow of Palestine belongs to the sub-genus Petronia.) In the house-sparrow (P. domesticus) the male differs much from the female and from the young. The young and the females are alike, and resemble 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, and by 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. That the males varied whilst young, and transmitted their variations to their offspring of both sexes, is the most probable. I may here add that I have, with little success, endeavoured, by consulting various works, to decide how far the period of variation in birds 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 (34. For instance, the males of Tanagra aestiva and Fringilla cyanea require three years, the male of Fringilla ciris four years, to complete their beautiful plumage. (See Audubon, ‘Ornith. Biography,’ vol. i. pp. 233, 280, 378). The Harlequin duck takes three years (ibid. vol. iii. p. 614). The male of the Gold pheasant, as I hear from Mr. Jenner Weir, can be distinguished from the female when about three months old, but he does not acquire his full splendour until the end of the September in the following year.), second, and fourth classes of cases; but they fail in the third, often in the fifth (35. Thus the Ibis tantalus and Grus americanus take four years, the Flamingo several years, and the Ardea ludovicana two years, before they acquire their perfect plumage. See Audubon, ibid. vol. i. p. 221; vol. iii. pp. 133, 139, 211.), and in the sixth small class. They apply, however, as far as I can judge, to a considerable majority of the species; and we must not forget the striking generalisation by Dr. W. Marshall with respect to the protuberances on the heads of birds. Whether or not the two rules generally hold good, we may conclude from the facts given in the eighth chapter, that the period of variation is 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 are first developed will falsely appear to us to be earlier than it really is. 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. (36. Mr. Blyth, in Charlesworth’s ‘Magazine of Natural History,’ vol. i. 1837, p. 300. Mr. Bartlett has informed me in regard to gold pheasants.) 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 power of reproduction is gained, it is a remarkable fact that various birds occasionally breed whilst retaining their immature plumage. (37. I have noticed the following cases in Audubon’s ‘Ornith. Biography.’ The redstart of America (Muscapica ruticilla, vol. i. p. 203). The Ibis tantalus takes four years to come to full maturity, but sometimes breeds in the second year (vol. iii. p. 133). The Grus americanus takes the same time, but breeds before acquiring its full plumage (vol. iii. p. 211). The adults of Ardea caerulea are blue, and the young white; and white, mottled, and mature blue birds may all be seen breeding together (vol. iv. p. 58): but Mr. Blyth informs me that certain herons apparently are dimorphic, for white and coloured individuals of the same age may be observed. The Harlequin duck (Anas histrionica, Linn.) takes three years to acquire its full plumage, though many birds breed in the second year (vol. iii. p. 614). The White-headed Eagle (Falco leucocephalus, vol. iii. p. 210) is likewise known to breed in its immature state. Some species of Oriolus (according to Mr. Blyth and Mr. Swinhoe, in ‘Ibis,’ July 1863, p. 68) likewise breed before they attain their full plumage.)

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, etc., 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. (38. See footnote 37 above.) 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 would prevail, which retained their immature dress for the longest period, and thus the character of the species would ultimately be modified. (39. Other animals, belonging to quite distinct classes, are either habitually or occasionally capable of breeding before they have fully acquired their adult characters. This is the case with the young males of the salmon. Several amphibians have been known to breed whilst retaining their larval structure. Fritz Müller has shewn (‘Facts and arguments for Darwin,’ Eng. trans. 1869, p. 79) that the males of several amphipod crustaceans become sexually mature whilst young; and I infer that this is a case of premature breeding, because they have not as yet acquired their fully-developed claspers. All such facts are highly interesting, as bearing on one means by which species may undergo great modifications of character.) If, on the other hand, the young never succeeded in obtaining a female, the habit of early reproduction would perhaps be sooner or later 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, with some of the birds of paradise, and with the crest and plumes of certain herons, for instance, the Ardea ludovicana. (40. Jerdon, ‘Birds of India,’ vol. iii. p. 507, on the peacock. Dr. Marshall thinks that the older and more brilliant males of birds of paradise, have an advantage over the younger males; see ‘Archives Neerlandaises,’ tom. vi. 1871.—On Ardea, Audubon, ibid. vol. iii. p. 139.) But it is doubtful whether the continued development of such feathers is the result of the selection of successive beneficial variations (though this is the most probable view with birds of paradise) 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 (41. For illustrative cases, see vol. iv. of Macgillivray’s ‘History of British Birds;’ on Tringa, etc., pp. 229, 271; on the Machetes, p. 172; on the Charadrius hiaticula, p. 118; on the Charadrius pluvialis, p. 94.), the young generally resemble the females,—as with the so-called goldfinch of North America, and apparently with the splendid Maluri of Australia. (42. For the goldfinch of N. America, Fringilla tristis, Linn., see Audubon, ‘Ornithological Biography,’ vol. i. p. 172. For the Maluri, Gould’s ‘Handbook of the Birds of Australia,’ vol. i. p. 318.) With those 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, and this is of much rarer occurrence, 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. (43. I am indebted to Mr. Blyth for information as to the Buphus; see also Jerdon, ‘Birds of India,’ vol. iii. p. 749. On the Anastomus, see Blyth, in ‘Ibis,’ 1867, p. 173.) 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. (44. On the Alca, see Macgillivray, ‘Hist. Brit. Birds,’ vol. v. p. 347. On the Fringilla leucophrys, Audubon, ibid. vol. ii. p. 89. I shall have hereafter to refer to the young of certain herons and egrets being white.) With respect to the third case, namely, that of the young having an intermediate character between the summer and winter adult plumages, Yarrell (45. ‘History of British Birds,’ vol. i. 1839, p. 159.) 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 females in their 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 by 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 it seems the most natural thing that the young should at first somewhat resemble the adults of the same sex, and gradually become more and more like them. 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; thus, the male blackbird (Turdus merula) can be distinguished in the nest from the female. 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 showing more pure white. (46. Audubon, ‘Ornith. Biography,’ vol. i. p. 113.) The males of a forest-thrush and of a rock-thrush (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. (47. Mr. C.A. Wright, in ‘Ibis,’ vol. vi. 1864, p. 65. Jerdon, ‘Birds of India,’ vol. i. p. 515. See also on the blackbird, Blyth in Charlesworth’s ‘Magazine of Natural History,’ vol. i. 1837, p. 113.) In the young blackbird the wing-feathers assume their mature character and become black after the others; on the other hand, in the two species just named the wing-feathers become blue before the others. The most probable view with reference to the cases in the present class 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 were first acquired; for, if the males had varied whilst quite young, their characters would probably have been transmitted to both sexes. (48. The following additional cases may be mentioned; the young males of Tanagra rubra can be distinguished from the young females (Audubon, ‘Ornith. Biography,’ vol. iv. p. 392), and so it is within the nestlings of a blue nuthatch, Dendrophila frontalis of India (Jerdon, ‘Birds of India,’ vol. i. p. 389). Mr. Blyth also informs me that the sexes of the stonechat, Saxicola rubicola, are distinguishable at a very early age. Mr. Salvin gives (‘Proc. Zoolog. Soc.’ 1870, p. 206) the case of a humming-bird, like the following one of Eustephanus.)

In Aithurus polytmus, a humming-bird, 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 chestnut-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 somewhat resemble 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 rendered beautiful independently; and not that 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 Rhynchaea 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 the females in number, 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 far exceeded the males, we could understand how the males at one time, and the females at another, 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 all six classes, that an intimate relation exists between the plumage of the young and the adults, either of one sex or both. 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 species of the same group, we do not know, but with respect to the form of transmission, one important determining cause seems to be the age at which the variations first appear.

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 of both are bright 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 bright-coloured species being more brilliant 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), of which the young are bright-coloured, and as these form old groups, we may infer that their early progenitors were likewise bright. With this exception, if we look to the birds of the world, it appears that their beauty has been much increased since that period, of which their immature plumage gives us a partial record.

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, so as to escape the notice of their enemies; or in some instances, so as to approach their prey unobserved, just as owls have had their plumage rendered soft, that their flight may not be overheard. Mr. Wallace remarks (49. ‘Westminster Review,’ July 1867, p. 5.) 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 with tree-haunting birds, strongly-pronounced colours have been acquired through sexual selection, but that a green tint has been acquired oftener than any other, from the additional advantage of protection.

In regard to birds which live on the ground, every one 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 night-jars when crouched on ground. Animals inhabiting deserts offer the most striking cases, for the bare surface affords no concealment, and nearly all the smaller quadrupeds, reptiles, and birds depend for safety on their colours. Mr. Tristram has remarked in regard to the inhabitants of the Sahara, that all are protected by their “isabelline or sand-colour.” (50. ‘Ibis,’ 1859, vol. i. p. 429, et seq. Dr. Rohlfs, however, remarks to me in a letter that according to his experience of the Sahara, this statement is too strong.) Calling to my recollection the desert-birds of South America, as well as most of the ground-birds of 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 their plumage coloured in a protective manner; and this colouring is all the more striking, as with most of these birds it differs 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 here 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 the two 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 by natural selection from inheriting the colours of their male parents; so that we must look to the law of sexually-limited transmission.

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 night-jar, 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.

Both sexes of many birds 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, blackcock, 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 much to their beauty; we see this in the bright yellow beak of the male blackbird, in the crimson skin over the eyes of the blackcock and capercailzie, in the brightly and variously coloured beak of the scoter-drake (Oidemia), in the red beak of the chough (Corvus graculus, Linn.), of the black swan, and the black stork. This leads me to remark that it is not 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. (51. No satisfactory explanation has ever been offered of the immense size, and still less of the bright colours, of the toucan’s beak. Mr. Bates (‘The Naturalist on the Amazons,’ vol. ii. 1863, p. 341) states that they use their beaks for reaching fruit at the extreme tips of the branches; and likewise, as stated by other authors, for extracting eggs and young birds from the nests of other birds. But, as Mr. Bates admits, the beak “can scarcely be considered a very perfectly-formed instrument for the end to which it is applied.” The great bulk of the beak, as shewn by its breadth, depth, as well as length, is not intelligible on the view, that it serves merely as an organ of prehension. Mr. Belt believes (‘The Naturalist in Nicaragua,’ p. 197) that the principal use of the beak is as a defence against enemies, especially to the female whilst nesting in a hole in a tree.) The naked skin, also, at the base of the beak and round the eyes is likewise often brilliantly coloured; and Mr. Gould, in speaking of one species (52. Rhamphastos carinatus, Gould’s ‘Monograph of Ramphastidae.’), 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 that toucans should be encumbered with immense beaks, though rendered as light as possible by their cancellated structure, for the display of fine colours (an object falsely appearing to us unimportant), 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, etc., 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, etc., have acquired their more or less completely white plumage through sexual selection. In some of these cases the plumage becomes white only at maturity. This is the case with certain gannets, tropic-birds, etc., 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 Anastomus oscitans, 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. (53. On Larus, Gavia, and Sterna, see Macgillivray, ‘History of British Birds,’ vol. v. pp. 515, 584, 626. On the Anser hyperboreus, Audubon, ‘Ornithological Biography,’ vol. iv. p. 562. On the Anastomus, Mr. Blyth, in ‘Ibis,’ 1867, p. 173.)

That aquatic birds have acquired a white plumage so much oftener 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. (54. It may be noticed that with vultures, which roam far and wide high in the air, like marine birds over the ocean, three or four species are almost wholly or largely white, and that many others are black. So that here again conspicuous colours may possibly aid the sexes in finding each other during the breeding-season.) 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 other 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 procure more food than the less strongly coloured individuals. Hence conspicuous colours cannot have been gradually acquired for this purpose through natural selection.

As sexual selection depends on so fluctuating an element as taste, we can understand how it is that, within the same group of birds having nearly the same habits, there should exist white or nearly white, as well as black, or nearly black species,—for instance, both white and black cockatoos, storks, ibises, swans, terns, and petrels. Piebald birds likewise sometimes occur in the same groups together with black and white species; 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, for the sexes often 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 the female.

It would even appear that mere novelty, or slight changes for the sake of change, have sometimes acted on female birds as a charm, like changes of fashion with us. Thus the males of some parrots can hardly be said to be more beautiful than the females, at least according to our taste, but they differ in such points, as in having a rose-coloured collar instead of “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. (55. See Jerdon on the genus Palaeornis, ‘Birds of India,’ vol. i. pp. 258-260.) As so many male birds have elongated tail-feathers or elongated crests for their chief ornament, the shortened tail, formerly described in the male of a humming-bird, and the shortened crest of the male goosander, seem like one of the many 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, as it appears, 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 in their winter plumage, of the allied Buphus coromandus 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 (56. The young of Ardea rufescens and A. caerulea of the United States are likewise white, the adults being coloured in accordance with their specific names. Audubon (‘Ornithological Biography,’ vol. iii. p. 416; vol. iv. p. 58) seems rather pleased at the thought that this remarkable change of plumage will greatly “disconcert the systematists.”), should for any special purpose have been 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 good reason to believe that whiteness has been gained by many birds as a sexual ornament. We may therefore conclude that some 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; and that the whiteness was afterwards retained by the young, whilst it was exchanged by the adults for more strongly-pronounced tints. But if we could look still further back 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, during a long line of descent, the adult progenitors of the Ardea asha, the Buphus, and of some allies, have undergone the following changes of colour: first, 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 its own sake.

Several writers have objected to the whole theory of sexual selection, by assuming that with animals and savages the taste of the female for certain colours or other ornaments would not remain constant for many generations; that first one colour and then another would be admired, and consequently that no permanent effect could be produced. We may admit that taste is fluctuating, but it is not quite arbitrary. It depends much on habit, as we see in mankind; and we may infer that this would hold good with birds and other animals. Even in our own dress, the general character lasts long, and the changes are to a certain extent graduated. Abundant evidence will be given in two places in a future chapter, that savages of many races have admired for many generations the same cicatrices on the skin, the same hideously perforated lips, nostrils, or ears, distorted heads, etc.; and these deformities present some analogy to the natural ornaments of various animals. Nevertheless, with savages such fashions do not endure for ever, as we may infer from the differences in this respect between allied tribes on the same continent. So again the raisers of fancy animals certainly have admired for many generations and still admire the same breeds; they earnestly desire slight changes, which are considered as improvements, but any great or sudden change is looked at as the greatest blemish. With birds in a state of nature we have no reason to suppose that they would admire an entirely new style of coloration, even if great and sudden variations often occurred, which is far from being the case. We know that dovecot pigeons do not willingly associate with the variously coloured fancy breeds; that albino birds do not commonly get partners in marriage; and that the black ravens of the Feroe Islands chase away their piebald brethren. But this dislike of a sudden change would not preclude their appreciating slight changes, any more than it does in the case of man. Hence with respect to taste, which depends on many elements, but partly on habit and partly on a love of novelty, there seems no improbability in animals admiring for a very long period the same general style of ornamentation or other attractions, and yet appreciating slight changes in colours, form, or sound.

A SUMMARY OF THE FOUR CHAPTERS ON BIRDS.

Most male birds are highly pugnacious during the breeding-season, and some possess weapons 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 giving forth strange cries, or 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 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 where the sexes differ, the males differ much more from one another 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 must determine the nature of the initial changes, and will have largely influenced the final result. The gradations, which may be observed between the males of allied species, indicate the nature of the steps through which they have passed. They explain also in the most interesting manner how certain characters have originated, such as the indented ocelli on the tail-feathers of the peacock, and the ball-and-socket ocelli on the wing-feathers of the Argus pheasant. It is evident that the brilliant colours, top-knots, fine plumes, etc., 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 in some cases produced a definite effect on both sexes, or sometimes on one sex alone, the more important result will have been an increased tendency to vary or to present more strongly-marked individual differences; and such differences will have afforded an excellent ground-work 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 have been transmitted sometimes in one way and sometimes in another, is not in most cases 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 more or less 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 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 species, in which the sexes differ in colour, it is possible or probable that some of the successive variations often tended to be transmitted equally to both sexes; but that when this occurred the females were prevented from acquiring the bright colours of the males, by the destruction which they suffered during incubation. There is no evidence that it is possible by natural selection to convert one form of transmission into another. But 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 were rendered as conspicuously coloured as the males, their instincts appear often to have been modified so that they were 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 brighter coloured than the males. They have, also, become so quarrelsome that they often fight together for the possession of the males, like the males of other pugnacious species for the possession of the females. If, as seems probable, such females habitually drive away their rivals, 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 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 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 if dangerous to the young, they will be eliminated through natural selection. Thus we can understand how it is that variations arising late in life have so often been preserved for the ornamentation 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. (57. I am greatly indebted to the kindness of Mr. Sclater for having looked over these four chapters on birds, and the two following ones on mammals. In this way I have been saved from making mistakes about the names of the species, and from stating anything as a fact which is known to this distinguished naturalist to be erroneous. But, of course, he is not at all answerable for the accuracy of the statements quoted by me from various authorities.)

XVII. SECONDARY SEXUAL CHARACTERS OF MAMMALS

The law of battle—Special weapons, confined to the males—Cause of absence of weapons in the female—Weapons common to both sexes, yet primarily acquired by the male—Other uses of such weapons—Their high importance—Greater size of the male—Means of defence—On the preference shown by either sex in the pairing of quadrupeds.

With mammals the male appears to win the female much more through the law of battle than through the display of his charms. The most timid animals, not provided with any special weapons for fighting, engage in desperate conflicts during the season of love. Two male hares have been seen to fight together until one was killed; male moles often fight, and sometimes with fatal results; male squirrels engage in frequent contests, “and often wound each other severely”; as do male beavers, so that “hardly a skin is without scars.” (1. See Waterton’s account of two hares fighting, ‘Zoologist,’ vol. i. 1843, p. 211. On moles, Bell, ‘Hist. of British Quadrupeds,’ 1st ed., p. 100. On squirrels, Audubon and Bachman, Viviparous Quadrupeds of N. America, 1846, p. 269. On beavers, Mr. A.H. Green, in ‘Journal of Linnean Society, Zoology,’ vol. x. 1869, p. 362.) I observed the same fact with the hides of the guanacoes in Patagonia; and on one occasion several were so absorbed in fighting that they fearlessly rushed close by me. Livingstone speaks of the males of the many animals in Southern Africa as almost invariably shewing the scars received in former contests.

The law of battle prevails with aquatic as with terrestrial mammals. It is notorious how desperately male seals fight, both with their teeth and claws, during the breeding-season; and their hides are likewise often covered with scars. Male sperm-whales are very jealous at this season; and in their battles “they often lock their jaws together, and turn on their sides and twist about”; so that their lower jaws often become distorted. (2. On the battles of seals, see Capt. C. Abbott in ‘Proc. Zool. Soc.’ 1868, p. 191; Mr. R. Brown, ibid. 1868, p. 436; also L. Lloyd, ‘Game Birds of Sweden,’ 1867, p. 412; also Pennant. On the sperm-whale see Mr. J.H. Thompson, in ‘Proc. Zool. Soc.’ 1867, p. 246.)

All male animals which are furnished with special weapons for fighting, are well known to engage in fierce battles. The courage and the desperate conflicts of stags have often been described; their skeletons have been found in various parts of the world, with the horns inextricably locked together, shewing how miserably the victor and vanquished had perished. (3. See Scrope (‘Art of Deer-stalking,’ p. 17) on the locking of the horns with the Cervus elaphus. Richardson, in ‘Fauna Bor. Americana,’ 1829, p. 252, says that the wapiti, moose, and reindeer have been found thus locked together. Sir A. Smith found at the Cape of Good Hope the skeletons of two gnus in the same condition.) No animal in the world is so dangerous as an elephant in must. Lord Tankerville has given me a graphic description of the battles between the wild bulls in Chillingham Park, the descendants, degenerated in size but not in courage, of the gigantic Bos primigenius. In 1861 several contended for mastery; and it was observed that two of the younger bulls attacked in concert the old leader of the herd, overthrew and disabled him, so that he was believed by the keepers to be lying mortally wounded in a neighbouring wood. But a few days afterwards one of the young bulls approached the wood alone; and then the “monarch of the chase,” who had been lashing himself up for vengeance, came out and, in a short time, killed his antagonist. He then quietly joined the herd, and long held undisputed sway. Admiral Sir B.J. Sulivan informs me that, when he lived in the Falkland Islands, he imported a young English stallion, which frequented the hills near Port William with eight mares. On these hills there were two wild stallions, each with a small troop of mares; “and it is certain that these stallions would never have approached each other without fighting. Both had tried singly to fight the English horse and drive away his mares, but had failed. One day they came in TOGETHER and attacked him. This was seen by the capitan who had charge of the horses, and who, on riding to the spot, found one of the two stallions engaged with the English horse, whilst the other was driving away the mares, and had already separated four from the rest. The capitan settled the matter by driving the whole party into the corral, for the wild stallions would not leave the mares.”

Male animals which are provided with efficient cutting or tearing teeth for the ordinary purposes of life, such as the carnivora, insectivora, and rodents, are seldom furnished with weapons especially adapted for fighting with their rivals. The case is very different with the males of many other animals. We see this in the horns of stags and of certain kinds of antelopes in which the females are hornless. With many animals the canine teeth in the upper or lower jaw, or in both, are much larger in the males than in the females, or are absent in the latter, with the exception sometimes of a hidden rudiment. Certain antelopes, the musk-deer, camel, horse, boar, various apes, seals, and the walrus, offer instances. In the females of the walrus the tusks are sometimes quite absent. (4. Mr. Lamont (‘Seasons with the Sea-Horses,’ 1861, p. 143) says that a good tusk of the male walrus weighs 4 pounds, and is longer than that of the female, which weighs about 3 pounds. The males are described as fighting ferociously. On the occasional absence of the tusks in the female, see Mr. R. Brown, ‘Proceedings, Zoological Society,’ 1868, p. 429.) In the male elephant of India and in the male dugong (5. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 283.) the upper incisors form offensive weapons. In the male narwhal the left canine alone is developed into the well-known, spirally-twisted, so-called horn, which is sometimes from nine to ten feet in length. It is believed that the males use these horns for fighting together; for “an unbroken one can rarely be got, and occasionally one may be found with the point of another jammed into the broken place.” (6. Mr. R. Brown, in ‘Proc. Zool. Soc.’ 1869, p. 553. See Prof. Turner, in ‘Journal of Anat. and Phys.’ 1872, p. 76, on the homological nature of these tusks. Also Mr. J.W. Clarke on two tusks being developed in the males, in ‘Proceedings of the Zoological Society,’ 1871, p. 42.) The tooth on the opposite side of the head in the male consists of a rudiment about ten inches in length, which is embedded in the jaw; but sometimes, though rarely, both are equally developed on the two sides. In the female both are always rudimentary. The male cachalot has a larger head than that of the female, and it no doubt aids him in his aquatic battles. Lastly, the adult male ornithorhynchus is provided with a remarkable apparatus, namely a spur on the foreleg, closely resembling the poison-fang of a venomous snake; but according to Harting, the secretion from the gland is not poisonous; and on the leg of the female there is a hollow, apparently for the reception of the spur. (7. Owen on the cachalot and Ornithorhynchus, ibid. vol. iii. pp. 638, 641. Harting is quoted by Dr. Zouteveen in the Dutch translation of this work, vol. ii. p. 292.)

When the males are provided with weapons which in the females are absent, there can be hardly a doubt that these serve for fighting with other males; and that they were acquired through sexual selection, and were transmitted to the male sex alone. It is not probable, at least in most cases, that the females have been prevented from acquiring such weapons, on account of their being useless, superfluous, or in some way injurious. On the contrary, as they are often used by the males for various purposes, more especially as a defence against their enemies, it is a surprising fact that they are so poorly developed, or quite absent, in the females of so many animals. With female deer the development during each recurrent season of great branching horns, and with female elephants the development of immense tusks, would be a great waste of vital power, supposing that they were of no use to the females. Consequently, they would have tended to be eliminated in the female through natural selection; that is, if the successive variations were limited in their transmission to the female sex, for otherwise the weapons of the males would have been injuriously affected, and this would have been a greater evil. On the whole, and from the consideration of the following facts, it seems probable that when the various weapons differ in the two sexes, this has generally depended on the kind of transmission which has prevailed.

As the reindeer is the one species in the whole family of Deer, in which the female is furnished with horns, though they are somewhat smaller, thinner, and less branched than in the male, it might naturally be thought that, at least in this case, they must be of some special service to her. The female retains her horns from the time when they are fully developed, namely, in September, throughout the winter until April or May, when she brings forth her young. Mr. Crotch made particular enquiries for me in Norway, and it appears that the females at this season conceal themselves for about a fortnight in order to bring forth their young, and then reappear, generally hornless. In Nova Scotia, however, as I hear from Mr. H. Reeks, the female sometimes retains her horns longer. The male on the other hand casts his horns much earlier, towards the end of November. As both sexes have the same requirements and follow the same habits of life, and as the male is destitute of horns during the winter, it is improbable that they can be of any special service to the female during this season, which includes the larger part of the time during which she is horned. Nor is it probable that she can have inherited horns from some ancient progenitor of the family of deer, for, from the fact of the females of so many species in all quarters of the globe not having horns, we may conclude that this was the primordial character of the group. (8. On the structure and shedding of the horns of the reindeer, Hoffberg, ‘Amoenitates Acad.’ vol. iv. 1788, p. 149. See Richardson, ‘Fauna Bor. Americana,’ p. 241, in regard to the American variety or species: also Major W. Ross King, ‘The Sportsman in Canada,’ 1866, p. 80.

The horns of the reindeer are developed at a most unusually early age; but what the cause of this may be is not known. The effect has apparently been the transference of the horns to both sexes. We should bear in mind that horns are always transmitted through the female, and that she has a latent capacity for their development, as we see in old or diseased females. (9. Isidore Geoffroy St.-Hilaire, ‘Essais de Zoolog. Générale,’ 1841, p. 513. Other masculine characters, besides the horns, are sometimes similarly transferred to the female; thus Mr. Boner, in speaking of an old female chamois (‘Chamois Hunting in the Mountains of Bavaria,’ 1860, 2nd ed., p. 363), says, “not only was the head very male-looking, but along the back there was a ridge of long hair, usually to be found only in bucks.”) Moreover the females of some other species of deer exhibit, either normally or occasionally, rudiments of horns; thus the female of Cervulus moschatus has “bristly tufts, ending in a knob, instead of a horn”; and “in most specimens of the female wapiti (Cervus canadensis) there is a sharp bony protuberance in the place of the horn.” (10. On the Cervulus, Dr. Gray, ‘Catalogue of Mammalia in the British Museum,’ part iii. p. 220. On the Cervus canadensis or wapiti, see Hon. J.D. Caton, ‘Ottawa Academy of Nat. Sciences,’ May 1868, p. 9.) From these several considerations we may conclude that the possession of fairly well-developed horns by the female reindeer, is due to the males having first acquired them as weapons for fighting with other males; and secondarily to their development from some unknown cause at an unusually early age in the males, and their consequent transference to both sexes.

Turning to the sheath-horned ruminants: with antelopes a graduated series can be formed, beginning with species, the females of which are completely destitute of horns—passing on to those which have horns so small as to be almost rudimentary (as with the Antilocapra americana, in which species they are present in only one out of four or five females (11. I am indebted to Dr. Canfield for this information; see also his paper in the ‘Proceedings of the Zoological Society,’ 1866, p. 105.))—to those which have fairly developed horns, but manifestly smaller and thinner than in the male and sometimes of a different shape (12. For instance the horns of the female Ant. euchore resemble those of a distinct species, viz. the Ant. dorcas var. Corine, see Desmarest, ‘Mammalogie,’ p. 455.),—and ending with those in which both sexes have horns of equal size. As with the reindeer, so with antelopes, there exists, as previously shewn, a relation between the period of the development of the horns and their transmission to one or both sexes; it is therefore probable that their presence or absence in the females of some species, and their more or less perfect condition in the females of other species, depends, not on their being of any special use, but simply on inheritance. It accords with this view that even in the same restricted genus both sexes of some species, and the males alone of others, are thus provided. It is also a remarkable fact that, although the females of Antilope bezoartica are normally destitute of horns, Mr. Blyth has seen no less than three females thus furnished; and there was no reason to suppose that they were old or diseased.

In all the wild species of goats and sheep the horns are larger in the male than in the female, and are sometimes quite absent in the latter. (13. Gray, ‘Catalogue of Mammalia, the British Museum,’ part iii. 1852, p. 160.) In several domestic breeds of these two animals, the males alone are furnished with horns; and in some breeds, for instance, in the sheep of North Wales, though both sexes are properly horned, the ewes are very liable to be hornless. I have been informed by a trustworthy witness, who purposely inspected a flock of these same sheep during the lambing season, that the horns at birth are generally more fully developed in the male than in the female. Mr. J. Peel crossed his Lonk sheep, both sexes of which always bear horns, with hornless Leicesters and hornless Shropshire Downs; and the result was that the male offspring had their horns considerably reduced, whilst the females were wholly destitute of them. These several facts indicate that, with sheep, the horns are a much less firmly fixed character in the females than in the males; and this leads us to look at the horns as properly of masculine origin.

With the adult musk-ox (Ovibos moschatus) the horns of the male are larger than those of the female, and in the latter the bases do not touch. (14. Richardson, ‘Fauna Bor. Americana,’ p. 278.) In regard to ordinary cattle Mr. Blyth remarks: “In most of the wild bovine animals the horns are both longer and thicker in the bull than in the cow, and in the cow-banteng (Bos sondaicus) the horns are remarkably small, and inclined much backwards. In the domestic races of cattle, both of the humped and humpless types, the horns are short and thick in the bull, longer and more slender in the cow and ox; and in the Indian buffalo, they are shorter and thicker in the bull, longer and more slender in the cow. In the wild gaour (B. gaurus) the horns are mostly both longer and thicker in the bull than in the cow.” (15. ‘Land and Water,’ 1867, p. 346.) Dr. Forsyth Major also informs me that a fossil skull, believed to be that of the female Bos etruscus, has been found in Val d’Arno, which is wholly without horns. In the Rhinoceros simus, as I may add, the horns of the female are generally longer but less powerful than in the male; and in some other species of rhinoceros they are said to be shorter in the female. (16. Sir Andrew Smith, ‘Zoology of S. Africa,’ pl. xix. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 624.) From these various facts we may infer as probable that horns of all kinds, even when they are equally developed in the two sexes, were primarily acquired by the male in order to conquer other males, and have been transferred more or less completely to the female.

The effects of castration deserve notice, as throwing light on this same point. Stags after the operation never renew their horns. The male reindeer, however, must be excepted, as after castration he does renew them. This fact, as well as the possession of horns by both sexes, seems at first to prove that the horns in this species do not constitute a sexual character (17. This is the conclusion of Seidlitz, ‘Die Darwinsche Theorie,’ 1871, p. 47.); but as they are developed at a very early age, before the sexes differ in constitution, it is not surprising that they should be unaffected by castration, even if they were aboriginally acquired by the male. With sheep both sexes properly bear horns; and I am informed that with Welch sheep the horns of the males are considerably reduced by castration; but the degree depends much on the age at which the operation is performed, as is likewise the case with other animals. Merino rams have large horns, whilst the ewes “generally speaking are without horns”; and in this breed castration seems to produce a somewhat greater effect, so that if performed at an early age the horns “remain almost undeveloped.” (18. I am much obliged to Prof. Victor Carus, for having made enquiries for me in Saxony on this subject. H. von Nathusius (‘Viehzucht,’ 1872, p. 64) says that the horns of sheep castrated at an early period, either altogether disappear or remain as mere rudiments; but I do not know whether he refers to merinos or to ordinary breeds.) On the Guinea coast there is a breed in which the females never bear horns, and, as Mr. Winwood Reade informs me, the rams after castration are quite destitute of them. With cattle, the horns of the males are much altered by castration; for instead of being short and thick, they become longer than those of the cow, but otherwise resemble them. The Antilope bezoartica offers a somewhat analogous case: the males have long straight spiral horns, nearly parallel to each other, and directed backwards; the females occasionally bear horns, but these when present are of a very different shape, for they are not spiral, and spreading widely, bend round with the points forwards. Now it

is a remarkable fact that, in the castrated male, as Mr. Blyth informs me, the horns are of the same peculiar shape as in the female, but longer and thicker. If we may judge from analogy, the female probably shews us, in these two cases of cattle and the antelope, the former condition of the horns in some early progenitor of each species. But why castration should lead to the reappearance of an early condition of the horns cannot be explained with any certainty. Nevertheless, it seems probable, that in nearly the same manner as the constitutional disturbance in the offspring, caused by a cross between two distinct species or races, often leads to the reappearance of long-lost characters (19. I have given various experiments and other evidence proving that this is the case, in my ‘Variation of Animals and Plants under Domestication,’ vol. ii. 1868, pp. 39-47.); so here, the disturbance in the constitution of the individual, resulting from castration, produces the same effect.

The tusks of the elephant, in the different species or races, differ according to sex, nearly as do the horns of ruminants. In India and Malacca the males alone are provided with well-developed tusks. The elephant of Ceylon is considered by most naturalists as a distinct race, but by some as a distinct species, and here “not one in a hundred is found with tusks, the few that possess them being exclusively males.” (20. Sir J. Emerson Tennent, ‘Ceylon,’ 1859, vol. ii. p. 274. For Malacca, ‘Journal of Indian Archipelago,’ vol. iv. p. 357.) The African elephant is undoubtedly distinct, and the female has large well-developed tusks, though not so large as those of the male.

These differences in the tusks of the several races and species of elephants—the great variability of the horns of deer, as notably in the wild reindeer—the occasional presence of horns in the female Antilope Bezoartica, and their frequent absence in the female of Antilocapra americana—the presence of two tusks in some few male narwhals—the complete absence of tusks in some female walruses—are all instances of the extreme variability of secondary sexual characters, and of their liability to differ in closely-allied forms.

Although tusks and horns appear in all cases to have been primarily developed as sexual weapons, they often serve other purposes. The elephant uses his tusks in attacking the tiger; according to Bruce, he scores the trunks of trees until they can be thrown down easily, and he likewise thus extracts the farinaceous cores of palms; in Africa he often uses one tusk, always the same, to probe the ground and thus ascertain whether it will bear his weight. The common bull defends the herd with his horns; and the elk in Sweden has been known, according to Lloyd, to strike a wolf dead with a single blow of his great horns. Many similar facts could be given. One of the most curious secondary uses to which the horns of an animal may be occasionally put is that observed by Captain Hutton (21. ‘Calcutta Journal of Natural History,’ vol. ii, 1843, p. 526.) with the wild goat (Capra aegagrus) of the Himalayas and, as it is also said with the ibex, namely that when the male accidentally falls from a height he bends inwards his head, and by alighting on his massive horns, breaks the shock. The female cannot thus use her horns, which are smaller, but from her more quiet disposition she does not need this strange kind of shield so much.

Each male animal uses his weapons in his own peculiar fashion. The common ram makes a charge and butts with such force with the bases of his horns, that I have seen a powerful man knocked over like a child. Goats and certain species of sheep, for instance the Ovis cycloceros of Afghanistan (22. Mr. Blyth, in ‘Land and Water,’ March, 1867, p. 134, on the authority of Capt. Hutton and others. For the wild Pembrokeshire goats, see the ‘Field,’ 1869, p. 150.), rear on their hind legs, and then not only butt, but “make a cut down and a jerk up, with the ribbed front of their scimitar-shaped horn, as with a sabre. When the O. cycloceros attacked a large domestic ram, who was a noted bruiser, he conquered him by the sheer novelty of his mode of fighting, always closing at once with his adversary, and catching him across the face and nose with a sharp drawing jerk of the head, and then bounding out of the way before the blow could be returned.” In Pembrokeshire a male goat, the master of a flock which during several generations had run wild, was known to have killed several males in single combat; this goat possessed enormous horns, measuring thirty-nine inches in a straight line from tip to tip. The common bull, as every one knows, gores and tosses his opponent; but the Italian buffalo is said never to use his horns: he gives a tremendous blow with his convex forehead, and then tramples on his fallen enemy with his knees—an instinct which the common bull does not possess. (23. M. E.M. Bailly, “Sur l’usage des cornes,” etc., .Annal des Sciences Nat.’ tom. ii. 1824, p. 369.) Hence a dog who pins a buffalo by the nose is immediately crushed. We must, however, remember that the Italian buffalo has been long domesticated, and it is by no means certain that the wild parent-form had similar horns. Mr. Bartlett informs me that when a female Cape buffalo (Bubalus caffer) was turned into an enclosure with a bull of the same species, she attacked him, and he in return pushed her about with great violence. But it was manifest to Mr. Bartlett that, had not the bull shewn dignified forbearance, he could easily have killed her by a single lateral thrust with his immense horns. The giraffe uses his short, hair-covered horns, which are rather longer in the male than in the female, in a curious manner; for, with his long neck, he swings his head to either side, almost upside down, with such force that I have seen a hard plank deeply indented by a single blow.

[Fig. 63. Oryx leucoryx, male (from the Knowsley Menagerie).]

With antelopes it is sometimes difficult to imagine how they can possibly use their curiously-shaped horns; thus the springboc (Ant. euchore) has rather short upright horns, with the sharp points bent inwards almost at right angles, so as to face each other; Mr. Bartlett does not know how they are used, but suggests that they would inflict a fearful wound down each side of the face of an antagonist. The slightly-curved horns of the Oryx leucoryx (Fig. 63) are directed backwards, and are of such length that their points reach beyond the middle of the back, over which they extend in almost parallel lines. Thus they seem singularly ill-fitted for fighting; but Mr. Bartlett informs me that when two of these animals prepare for battle, they kneel down, with their heads between their fore legs, and in this attitude the horns stand nearly parallel and close to the ground, with the points directed forwards and a little upwards. The combatants then gradually approach each other, and each endeavours to get the upturned points under the body of the other; if one succeeds in doing this, he suddenly springs up, throwing up his head at the same time, and can thus wound or perhaps even transfix his antagonist. Both animals always kneel down, so as to guard as far as possible against this manoeuvre. It has been recorded that one of these antelopes has used his horn with effect even against a lion; yet from being forced to place his head between the forelegs in order to bring the points of the horns forward, he would generally be under a great disadvantage when attacked by any other animal. It is, therefore, not probable that the horns have been modified into their present great length and peculiar position, as a protection against beasts of prey. We can however see that, as soon as some ancient male progenitor of the Oryx acquired moderately long horns, directed a little backwards, he would be compelled, in his battles with rival males, to bend his head somewhat inwards or downwards, as is now done by certain stags; and it is not improbable that he might have acquired the habit of at first occasionally and afterwards of regularly kneeling down. In this case it is almost certain that the males which possessed the longest horns would have had a great advantage over others with shorter horns; and then the horns would gradually have been rendered longer and longer, through sexual selection, until they acquired their present extraordinary length and position.

With stags of many kinds the branches of the horns offer a curious case of difficulty; for certainly a single straight point would inflict a much more serious wound than several diverging ones. In Sir Philip Egerton’s museum there is a horn of the red-deer (Cervus elaphus), thirty inches in length, with “not fewer than fifteen snags or branches”; and at Moritzburg there is still preserved a pair of antlers of a red-deer, shot in 1699 by Frederick I., one of which bears the astonishing number of thirty-three branches and the other twenty-seven, making altogether sixty branches. Richardson figures a pair of antlers of the wild reindeer with twenty-nine points. (24. On the horns of red-deer, Owen, ‘British Fossil Mammals,’ 1846, p. 478; Richardson on the horns of the reindeer, ‘Fauna Bor. Americana,’ 1829, p. 240. I am indebted to Prof. Victor Carus, for the Moritzburg case.) From the manner in which the horns are branched, and more especially from deer being known occasionally to fight together by kicking with their fore-feet (25. Hon. J.D. Caton (‘Ottawa Acad. of Nat. Science,’ May 1868, p. 9) says that the American deer fight with their fore-feet, after “the question of superiority has been once settled and acknowledged in the herd.” Bailly, ‘Sur l’Usage des cornes,’ ‘Annales des Sciences Nat.’ tom. ii. 1824, p. 371.), M. Bailly actually comes to the conclusion that their horns are more injurious than useful to them. But this author overlooks the pitched battles between rival males. As I felt much perplexed about the use or advantage of the branches, I applied to Mr. McNeill of Colonsay, who has long and carefully observed the habits of red-deer, and he informs me that he has never seen some of the branches brought into use, but that the brow antlers, from inclining downwards, are a great protection to the forehead, and their points are likewise used in attack. Sir Philip Egerton also informs me both as to red-deer and fallow-deer that, in fighting, they suddenly dash together, and getting their horns fixed against each other’s bodies, a desperate struggle ensues. When one is at last forced to yield and turn round, the victor endeavours to plunge his brow antlers into his defeated foe. It thus appears that the upper branches are used chiefly or exclusively for pushing and fencing. Nevertheless in some species the upper branches are used as weapons of offence; when a man was attacked by a wapiti deer (Cervus canadensis) in Judge Caton’s park in Ottawa, and several men tried to rescue him, the stag “never raised his head from the ground; in fact he kept his face almost flat on the ground, with his nose nearly between his fore feet, except when he rolled his head to one side to take a new observation preparatory to a plunge.” In this position the ends of the horns were directed against his adversaries. “In rolling his head he necessarily raised it somewhat, because his antlers were so long that he could not roll his head without raising them on one side, while, on the other side they touched the ground.” The stag by this procedure gradually drove the party of rescuers backwards to a distance of 150 or 200 feet; and the attacked man was killed. (26. See a most interesting account in the Appendix to Hon. J.D. Caton’s paper, as above quoted.)

[Fig. 64. Strepsiceros Kudu (from Sir Andrew Smith’s ‘Zoology of South Africa.’]

Although the horns of stags are efficient weapons, there can, I think, be no doubt that a single point would have been much more dangerous than a branched antler; and Judge Caton, who has had large experience with deer, fully concurs in this conclusion. Nor do the branching horns, though highly important as a means of defence against rival stags, appear perfectly well adapted for this purpose, as they are liable to become interlocked. The suspicion has therefore crossed my mind that they may serve in part as ornaments. That the branched antlers of stags as well as the elegant lyrated horns of certain antelopes, with their graceful double curvature (Fig. 64), are ornamental in our eyes, no one will dispute. If, then, the horns, like the splendid accoutrements of the knights of old, add to the noble appearance of stags and antelopes, they may have been modified partly for this purpose, though mainly for actual service in battle; but I have no evidence in favour of this belief.

An interesting case has lately been published, from which it appears that the horns of a deer in one district in the United States are now being modified through sexual and natural selection. A writer in an excellent American Journal (27. The ‘American Naturalist,’ Dec. 1869, p. 552.) says, that he has hunted for the last twenty-one years in the Adirondacks, where the Cervus virginianus abounds. About fourteen years ago he first heard of SPIKE-HORN BUCKS. These became from year to year more common; about five years ago he shot one, and afterwards another, and now they are frequently killed. “The spike-horn differs greatly from the common antler of the C. virginianus. It consists of a single spike, more slender than the antler, and scarcely half so long, projecting forward from the brow, and terminating in a very sharp point. It gives a considerable advantage to its possessor over the common buck. Besides enabling him to run more swiftly through the thick woods and underbrush (every hunter knows that does and yearling bucks run much more rapidly than the large bucks when armed with their cumbrous antlers), the spike-horn is a more effective weapon than the common antler. With this advantage the spike-horn bucks are gaining upon the common bucks, and may, in time, entirely supersede them in the Adirondacks. Undoubtedly, the first spike-horn buck was merely an accidental freak of nature. But his spike-horns gave him an advantage, and enabled him to propagate his peculiarity. His descendants having a like advantage, have propagated the peculiarity in a constantly increasing ratio, till they are slowly crowding the antlered deer from the region they inhabit.” A critic has well objected to this account by asking, why, if the simple horns are now so advantageous, were the branched antlers of the parent-form ever developed? To this I can only answer by remarking, that a new mode of attack with new weapons might be a great advantage, as shewn by the case of the Ovis cycloceros, who thus conquered a domestic ram famous for his fighting power. Though the branched antlers of a stag are well adapted for fighting with his rivals, and though it might be an advantage to the prong-horned variety slowly to acquire long and branched horns, if he had to fight only with others of the same kind, yet it by no means follows that branched horns would be the best fitted for conquering a foe differently armed. In the foregoing case of the Oryx leucoryx, it is almost certain that the victory would rest with an antelope having short horns, and who therefore did not need to kneel down, though an oryx might profit by having still longer horns, if he fought only with his proper rivals.

Male quadrupeds, which are furnished with tusks, use them in various ways, as in the case of horns. The boar strikes laterally and upwards; the musk-deer downwards with serious effect. (28. Pallas, ‘Spicilegia Zoologica,’ fasc. xiii. 1779, p. 18.) The walrus, though having so short a neck and so unwieldy a body, “can strike either upwards, or downwards, or sideways, with equal dexterity.” (29. Lamont, ‘Seasons with the Sea-Horses,’ 1861, p. 141.) I was informed by the late Dr. Falconer, that the Indian elephant fights in a different manner according to the position and curvature of his tusks. When they are directed forwards and upwards he is able to fling a tiger to a great distance—it is said to even thirty feet; when they are short and turned downwards he endeavours suddenly to pin the tiger to the ground and, in consequence, is dangerous to the rider, who is liable to be jerked off the howdah. (30. See also Corse (‘Philosophical Transactions,’ 1799, p. 212) on the manner in which the short-tusked Mooknah variety attacks other elephants.)

Very few male quadrupeds possess weapons of two distinct kinds specially adapted for fighting with rival males. The male muntjac-deer (Cervulus), however, offers an exception, as he is provided with horns and exserted canine teeth. But we may infer from what follows that one form of weapon has often been replaced in the course of ages by another. With ruminants the development of horns generally stands in an inverse relation with that of even moderately developed canine teeth. Thus camels, guanacoes, chevrotains, and musk-deer, are hornless, and they have efficient canines; these teeth being “always of smaller size in the females than in the males.” The Camelidae have, in addition to their true canines, a pair of canine-shaped incisors in their upper jaws. (31. Owen, ‘Anatomy of Vertebrates,’ vol. iii. p. 349.) Male deer and antelopes, on the other hand, possess horns, and they rarely have canine teeth; and these, when present, are always of small size, so that it is doubtful whether they are of any service in their battles. In Antilope montana they exist only as rudiments in the young male, disappearing as he grows old; and they are absent in the female at all ages; but the females of certain other antelopes and of certain deer have been known occasionally to exhibit rudiments of these teeth. (32. See Ruppell (in ‘Proc. Zoolog. Soc.’ Jan. 12, 1836, p. 3) on the canines in deer and antelopes, with a note by Mr. Martin on a female American deer. See also Falconer (‘Palaeont. Memoirs and Notes,’ vol. i. 1868, p. 576) on canines in an adult female deer. In old males of the musk-deer the canines (Pallas, ‘Spic. Zoolog.’ fasc. xiii. 1779, p. 18) sometimes grow to the length of three inches, whilst in old females a rudiment projects scarcely half an inch above the gums.) Stallions have small canine teeth, which are either quite absent or rudimentary in the mare; but they do not appear to be used in fighting, for stallions bite with their incisors, and do not open their mouths wide like camels and guanacoes. Whenever the adult male possesses canines, now inefficient, whilst the female has either none or mere rudiments, we may conclude that the early male progenitor of the species was provided with efficient canines, which have been partially transferred to the females. The reduction of these teeth in the males seems to have followed from some change in their manner of fighting, often (but not in the horse) caused by the development of new weapons.

Tusks and horns are manifestly of high importance to their possessors, for their development consumes much organised matter. A single tusk of the Asiatic elephant—one of the extinct woolly species—and of the African elephant, have been known to weigh respectively 150, 160, and 180 pounds; and even greater weights have been given by some authors. (33. Emerson Tennent, ‘Ceylon,’ 1859, vol. ii. p. 275; Owen, ‘British Fossil Mammals,’ 1846, p. 245.) With deer, in which the horns are periodically renewed, the drain on the constitution must be greater; the horns, for instance, of the moose weigh from fifty to sixty pounds, and those of the extinct Irish elk from sixty to seventy pounds—the skull of the latter weighing on an average only five pounds and a quarter. Although the horns are not periodically renewed in sheep, yet their development, in the opinion of many agriculturists, entails a sensible loss to the breeder. Stags, moreover, in escaping from beasts of prey are loaded with an additional weight for the race, and are greatly retarded in passing through a woody country. The moose, for instance, with horns extending five and a half feet from tip to tip, although so skilful in their use that he will not touch or break a twig when walking quietly, cannot act so dexterously whilst rushing away from a pack of wolves. “During his progress he holds his nose up, so as to lay the horns horizontally back; and in this attitude cannot see the ground distinctly.” (34. Richardson, ‘Fauna Bor. Americana,’ on the moose, Alces palmata, pp. 236, 237; on the expanse of the horns, ‘Land and Water,’ 1869, p. 143. See also Owen, ‘British Fossil Mammals,’ on the Irish elk, pp. 447, 455.) The tips of the horns of the great Irish elk were actually eight feet apart! Whilst the horns are covered with velvet, which lasts with red-deer for about twelve weeks, they are extremely sensitive to a blow; so that in Germany the stags at this time somewhat change their habits, and avoiding dense forests, frequent young woods and low thickets. (35. ‘Forest Creatures,’ by C. Boner, 1861, p. 60.) These facts remind us that male birds have acquired ornamental plumes at the cost of retarded flight, and other ornaments at the cost of some loss of power in their battles with rival males.

With mammals, when, as is often the case, the sexes differ in size, the males are almost always larger and stronger. I am informed by Mr. Gould that this holds good in a marked manner with the marsupials of Australia, the males of which appear to continue growing until an unusually late age. But the most extraordinary case is that of one of the seals (Callorhinus ursinus), a full-grown female weighing less than one-sixth of a full-grown male. (36. See the very interesting paper by Mr. J.A. Allen in ‘Bull. Mus. Comp. Zoology of Cambridge, United States,’ vol. ii. No. 1, p. 82. The weights were ascertained by a careful observer, Capt. Bryant. Dr. Gill in ‘The American Naturalist,’ January, 1871, Prof. Shaler on the relative size of the sexes of whales, ‘American Naturalist,’ January, 1873.) Dr. Gill remarks that it is with the polygamous seals, the males of which are well known to fight savagely together, that the sexes differ much in size; the monogamous species differing but little. Whales also afford evidence of the relation existing between the pugnacity of the males and their large size compared with that of the female; the males of the right-whales do not fight together, and they are not larger, but rather smaller, than their females; on the other hand, male sperm-whales fight much together, and their bodies are “often found scarred with the imprint of their rival’s teeth,” and they are double the size of the females. The greater strength of the male, as Hunter long ago remarked (37. ‘Animal Economy,’ p. 45.), is invariably displayed in those parts of the body which are brought into action in fighting with rival males—for instance, in the massive neck of the bull. Male quadrupeds are also more courageous and pugnacious than the females. There can be little doubt that these characters have been gained, partly through sexual selection, owing to a long series of victories, by the stronger and more courageous males over the weaker, and partly through the inherited effects of use. It is probable that the successive variations in strength, size, and courage, whether due to mere variability or to the effects of use, by the accumulation of which male quadrupeds have acquired these characteristic qualities, occurred rather late in life, and were consequently to a large extent limited in their transmission to the same sex.

From these considerations I was anxious to obtain information as to the Scotch deer-hound, the sexes of which differ more in size than those of any other breed (though blood-hounds differ considerably), or than in any wild canine species known to me. Accordingly, I applied to Mr. Cupples, well known for his success with this breed, who has weighed and measured many of his own dogs, and who has with great kindness collected for me the following facts from various sources. Fine male dogs, measured at the shoulder, range from 28 inches, which is low, to 33 or even 34 inches in height; and in weight from 80 pounds, which is light, to 120 pounds, or even more. The females range in height from 23 to 27, or even to 28 inches; and in weight from 50 to 70, or even 80 pounds. (38. See also Richardson’s ‘Manual on the Dog,’ p. 59. Much valuable information on the Scottish deer-hound is given by Mr. McNeill, who first called attention to the inequality in size between the sexes, in Scrope’s ‘Art of Deer-Stalking.’ I hope that Mr. Cupples will keep to his intention of publishing a full account and history of this famous breed.) Mr. Cupples concludes that from 95 to 100 pounds for the male, and 70 for the female, would be a safe average; but there is reason to believe that formerly both sexes attained a greater weight. Mr. Cupples has weighed puppies when a fortnight old; in one litter the average weight of four males exceeded that of two females by six and a half ounces; in another litter the average weight of four males exceeded that of one female by less than one ounce; the same males when three weeks old, exceeded the female by seven and a half ounces, and at the age of six weeks by nearly fourteen ounces. Mr. Wright of Yeldersley House, in a letter to Mr. Cupples, says: “I have taken notes on the sizes and weights of puppies of many litters, and as far as my experience goes, dog-puppies as a rule differ very little from bitches till they arrive at about five or six months old; and then the dogs begin to increase, gaining upon the bitches both in weight and size. At birth, and for several weeks afterwards, a bitch-puppy will occasionally be larger than any of the dogs, but they are invariably beaten by them later.” Mr. McNeill, of Colonsay, concludes that “the males do not attain their full growth till over two years old, though the females attain it sooner.” According to Mr. Cupples’ experience, male dogs go on growing in stature till they are from twelve to eighteen months old, and in weight till from eighteen to twenty-four months old; whilst the females cease increasing in stature at the age of from nine to fourteen or fifteen months, and in weight at the age of from twelve to fifteen months. From these various statements it is clear that the full difference in size between the male and female Scotch deer-hound is not acquired until rather late in life. The males almost exclusively are used for coursing, for, as Mr. McNeill informs me, the females have not sufficient strength and weight to pull down a full-grown deer. From the names used in old legends, it appears, as I hear from Mr. Cupples, that, at a very ancient period, the males were the most celebrated, the females being mentioned only as the mothers of famous dogs. Hence, during many generations, it is the male which has been chiefly tested for strength, size, speed, and courage, and the best will have been bred from. As, however, the males do not attain their full dimensions until rather late in life, they will have tended, in accordance with the law often indicated, to transmit their characters to their male offspring alone; and thus the great inequality in size between the sexes of the Scotch deer-hound may probably be accounted for.

[Fig. 65. Head of Common wild boar, in prime of life (from Brehm).]

The males of some few quadrupeds possess organs or parts developed solely as a means of defence against the attacks of other males. Some kinds of deer use, as we have seen, the upper branches of their horns chiefly or exclusively for defending themselves; and the Oryx antelope, as I am informed by Mr. Bartlett, fences most skilfully with his long, gently curved horns; but these are likewise used as organs of offence. The same observer remarks that rhinoceroses in fighting, parry each other’s sidelong blows with their horns, which clatter loudly together, as do the tusks of boars. Although wild boars fight desperately, they seldom, according to Brehm, receive fatal wounds, as the blows fall on each other’s tusks, or on the layer of gristly skin covering the shoulder, called by the German hunters, the shield; and here we have a part specially modified for defence. With boars in the prime of life (Fig. 65) the tusks in the lower jaw are used for fighting, but they become in old age, as Brehm states, so much curved inwards and upwards over the snout that they can no longer be used in this way. They may, however, still serve, and even more effectively, as a means of defence. In compensation for the loss of the lower tusks as weapons of offence, those in the upper jaw, which always project a little laterally, increase in old age so much in length and curve so much upwards that they can be used for attack. Nevertheless, an old boar is not so dangerous to man as one at the age of six or seven years. (39. Brehm, ‘Thierleben,’ B. ii. ss. 729-732.)

[Fig. 66. Skull of the Babirusa Pig (from Wallace’s ‘Malay Archipelago’).]

In the full-grown male Babirusa pig of Celebes (Fig. 66), the lower tusks are formidable weapons, like those of the European boar in the prime of life, whilst the upper tusks are so long and have their points so much curled inwards, sometimes even touching the forehead, that they are utterly useless as weapons of attack. They more nearly resemble horns than teeth, and are so manifestly useless as teeth that the animal was formerly supposed to rest his head by hooking them on to a branch! Their convex surfaces, however, if the head were held a little laterally, would serve as an excellent guard; and hence, perhaps, it is that in old animals they “are generally broken off, as if by fighting.” (40. See Mr. Wallace’s interesting account of this animal, ‘The Malay Archipelago,’ 1869, vol. i. p. 435.) Here, then, we have the curious case of the upper tusks of the Babirusa regularly assuming during the prime of life a structure which apparently renders them fitted only for defence; whilst in the European boar the lower tusks assume in a less degree and only during old age nearly the same form, and then serve in like manner solely for defence.

[Fig. 67. Head of female Aethiopian wart-hog, from ‘Proc. Zool. Soc.’ 1869, shewing the same characters as the male, though on a reduced scale. N.B. When the engraving was first made, I was under the impression that it represented the male.]

In the wart-hog (see Phacochoerus aethiopicus, Fig. 67) the tusks in the upper jaw of the male curve upwards during the prime of life, and from being pointed serve as formidable weapons. The tusks in the lower jaw are sharper than those in the upper, but from their shortness it seems hardly possible that they can be used as weapons of attack. They must, however, greatly strengthen those in the upper jaw, from being ground so as to fit closely against their bases. Neither the upper nor the lower tusks appear to have been specially modified to act as guards, though no doubt they are to a certain extent used for this purpose. But the wart-hog is not destitute of other special means of protection, for it has, on each side of the face, beneath the eyes, a rather stiff, yet flexible, cartilaginous, oblong pad (Fig. 67), which projects two or three inches outwards; and it appeared to Mr. Bartlett and myself, when viewing the living animal, that these pads, when struck from beneath by the tusks of an opponent, would be turned upwards, and would thus admirably protect the somewhat prominent eyes. I may add, on the authority of Mr. Bartlett, that these boars when fighting stand directly face to face.

Lastly, the African river-hog (Potomochoerus penicillatus) has a hard cartilaginous knob on each side of the face beneath the eyes, which answers to the flexible pad of the wart-hog; it has also two bony prominences on the upper jaw above the nostrils. A boar of this species in the Zoological Gardens recently broke into the cage of the wart-hog. They fought all night long, and were found in the morning much exhausted, but not seriously wounded. It is a significant fact, as shewing the purposes of the above-described projections and excrescences, that these were covered with blood, and were scored and abraded in an extraordinary manner.

Although the males of so many members of the pig family are provided with weapons, and as we have just seen with means of defence, these weapons seem to have been acquired within a rather late geological period. Dr. Forsyth Major specifies (41. ‘Atti della Soc. Italiana di Sc. Nat.’ 1873, vol. xv. fasc. iv.) several miocene species, in none of which do the tusks appear to have been largely developed in the males; and Professor Rutimeyer was formerly struck with this same fact.

The mane of the lion forms a good defence against the attacks of rival lions, the one danger to which he is liable; for the males, as Sir A. Smith informs me, engage in terrible battles, and a young lion dares not approach an old one. In 1857 a tiger at Bromwich broke into the cage of a lion and a fearful scene ensued: “the lion’s mane saved his neck and head from being much injured, but the tiger at last succeeded in ripping up his belly, and in a few minutes he was dead.” (42. ‘The Times,’ Nov. 10, 1857. In regard to the Canada lynx, see Audubon and Bachman, ‘Quadrupeds of North America,’ 1846, p. 139.) The broad ruff round the throat and chin of the Canadian lynx (Felis canadensis) is much longer in the male than in the female; but whether it serves as a defence I do not know. Male seals are well known to fight desperately together, and the males of certain kinds (Otaria jubata) (43. Dr. Murie, on Otaria, ‘Proc. Zoolog. Soc.’ 1869, p. 109. Mr. J.A. Allen, in the paper above quoted (p. 75), doubts whether the hair, which is longer on the neck in the male than in the female, deserves to be called a mane.) have great manes, whilst the females have small ones or none. The male baboon of the Cape of Good Hope (Cynocephalus porcarius) has a much longer mane and larger canine teeth than the female; and the mane probably serves as a protection, for, on asking the keepers in the Zoological Gardens, without giving them any clue to my object, whether any of the monkeys especially attacked each other by the nape of the neck, I was answered that this was not the case, except with the above baboon. In the Hamadryas baboon, Ehrenberg compares the mane of the adult male to that of a young lion, whilst in the young of both sexes and in the female the mane is almost absent.

It appeared to me probable that the immense woolly mane of the male American bison, which reaches almost to the ground, and is much more developed in the males than in the females, served as a protection to them in their terrible battles; but an experienced hunter told Judge Caton that he had never observed anything which favoured this belief. The stallion has a thicker and fuller mane than the mare; and I have made particular inquiries of two great trainers and breeders, who have had charge of many entire horses, and am assured that they “invariably endeavour to seize one another by the neck.” It does not, however, follow from the foregoing statements, that when the hair on the neck serves as a defence, that it was originally developed for this purpose, though this is probable in some cases, as in that of the lion. I am informed by Mr. McNeill that the long hairs on the throat of the stag (Cervus elaphus) serve as a great protection to him when hunted, for the dogs generally endeavour to seize him by the throat; but it is not probable that these hairs were specially developed for this purpose; otherwise the young and the females would have been equally protected.

CHOICE IN PAIRING BY EITHER SEX OF QUADRUPEDS.

Before describing in the next chapter, the differences between the sexes in voice, odours emitted, and ornaments, it will be convenient here to consider whether the sexes exert any choice in their unions. Does the female prefer any particular male, either before or after the males may have fought together for supremacy; or does the male, when not a polygamist, select any particular female? The general impression amongst breeders seems to be that the male accepts any female; and this owing to his eagerness, is, in most cases, probably the truth. Whether the female as a general rule indifferently accepts any male is much more doubtful. In the fourteenth chapter, on Birds, a considerable body of direct and indirect evidence was advanced, shewing that the female selects her partner; and it would be a strange anomaly if female quadrupeds, which stand higher in the scale and have higher mental powers, did not generally, or at least often, exert some choice. The female could in most cases escape, if wooed by a male that did not please or excite her; and when pursued by several males, as commonly occurs, she would often have the opportunity, whilst they were fighting together, of escaping with some one male, or at least of temporarily pairing with him. This latter contingency has often been observed in Scotland with female red-deer, as I am informed by Sir Philip Egerton and others. (44. Mr. Boner, in his excellent description of the habits of the red-deer in Germany (‘Forest Creatures,’ 1861, p. 81) says, “while the stag is defending his rights against one intruder, another invades the sanctuary of his harem, and carries off trophy after trophy.” Exactly the same thing occurs with seals; see Mr. J.A. Allen, ibid. p. 100.)

It is scarcely possible that much should be known about female quadrupeds in a state of nature making any choice in their marriage unions. The following curious details on the courtship of one of the eared seals (Callorhinus ursinus) are given (45. Mr. J.A. Allen in ‘Bull. Mus. Comp. Zoolog. of Cambridge, United States,’ vol. ii. No. 1, p. 99.) on the authority of Capt. Bryant, who had ample opportunities for observation. He says, “Many of the females on their arrival at the island where they breed appear desirous of returning to some particular male, and frequently climb the outlying rocks to overlook the rookeries, calling out and listening as if for a familiar voice. Then changing to another place they do the same again...As soon as a female reaches the shore, the nearest male goes down to meet her, making meanwhile a noise like the clucking of a hen to her chickens. He bows to her and coaxes her until he gets between her and the water so that she cannot escape him. Then his manner changes, and with a harsh growl he drives her to a place in his harem. This continues until the lower row of harems is nearly full. Then the males higher up select the time when their more fortunate neighbours are off their guard to steal their wives. This they do by taking them in their mouths and lifting them over the heads of the other females, and carefully placing them in their own harem, carrying them as cats do their kittens. Those still higher up pursue the same method until the whole space is occupied. Frequently a struggle ensues between two males for the possession of the same female, and both seizing her at once pull her in two or terribly lacerate her with their teeth. When the space is all filled, the old male walks around complacently reviewing his family, scolding those who crowd or disturb the others, and fiercely driving off all intruders. This surveillance always keeps him actively occupied.”

As so little is known about the courtship of animals in a state of nature, I have endeavoured to discover how far our domesticated quadrupeds evince any choice in their unions. Dogs offer the best opportunity for observation, as they are carefully attended to and well understood. Many breeders have expressed a strong opinion on this head. Thus, Mr. Mayhew remarks, “The females are able to bestow their affections; and tender recollections are as potent over them as they are known to be in other cases, where higher animals are concerned. Bitches are not always prudent in their loves, but are apt to fling themselves away on curs of low degree. If reared with a companion of vulgar appearance, there often springs up between the pair a devotion which no time can afterwards subdue. The passion, for such it really is, becomes of a more than romantic endurance.” Mr. Mayhew, who attended chiefly to the smaller breeds, is convinced that the females are strongly attracted by males of a large size. (46. ‘Dogs: their Management,’ by E. Mayhew, M.R.C.V.S., 2nd ed., 1864, pp. 187-192.) The well-known veterinary Blaine states (47. Quoted by Alex. Walker, ‘On Intermarriage,’ 1838, p. 276; see also p. 244.) that his own female pug dog became so attached to a spaniel, and a female setter to a cur, that in neither case would they pair with a dog of their own breed until several weeks had elapsed. Two similar and trustworthy accounts have been given me in regard to a female retriever and a spaniel, both of which became enamoured with terrier-dogs.

Mr. Cupples informs me that he can personally vouch for the accuracy of the following more remarkable case, in which a valuable and wonderfully-intelligent female terrier loved a retriever belonging to a neighbour to such a degree, that she had often to be dragged away from him. After their permanent separation, although repeatedly shewing milk in her teats, she would never acknowledge the courtship of any other dog, and to the regret of her owner never bore puppies. Mr. Cupples also states, that in 1868, a female deerhound in his kennel thrice produced puppies, and on each occasion shewed a marked preference for one of the largest and handsomest, but not the most eager, of four deerhounds living with her, all in the prime of life. Mr. Cupples has observed that the female generally favours a dog whom she has associated with and knows; her shyness and timidity at first incline her against a strange dog. The male, on the contrary, seems rather inclined towards strange females. It appears to be rare when the male refuses any particular female, but Mr. Wright, of Yeldersley House, a great breeder of dogs, informs me that he has known some instances; he cites the case of one of his own deerhounds, who would not take any notice of a particular female mastiff, so that another deerhound had to be employed. It would be superfluous to give, as I could, other instances, and I will only add that Mr. Barr, who has carefully bred many bloodhounds, states that in almost every instance particular individuals of opposite sexes shew a decided preference for each other. Finally, Mr. Cupples, after attending to this subject for another year, has written to me, “I have had full confirmation of my former statement, that dogs in breeding form decided preferences for each other, being often influenced by size, bright colour, and individual characters, as well as by the degree of their previous familiarity.”

In regard to horses, Mr. Blenkiron, the greatest breeder of race-horses in the world, informs me that stallions are so frequently capricious in their choice, rejecting one mare and without any apparent cause taking to another, that various artifices have to be habitually used. The famous Monarque, for instance, would never consciously look at the dam of Gladiateur, and a trick had to be practised. We can partly see the reason why valuable race-horse stallions, which are in such demand as to be exhausted, should be so particular in their choice. Mr. Blenkiron has never known a mare reject a horse; but this has occurred in Mr. Wright’s stable, so that the mare had to be cheated. Prosper Lucas (48. ‘Traité de l’Héréd. Nat.’ tom. ii. 1850, p. 296.) quotes various statements from French authorities, and remarks, “On voit des étalons qui s’eprennent d’une jument, et negligent toutes les autres.” He gives, on the authority of Baelen, similar facts in regard to bulls; and Mr. H. Reeks assures me that a famous short-horn bull belonging to his father “invariably refused to be matched with a black cow.” Hoffberg, in describing the domesticated reindeer of Lapland says, “Foeminae majores et fortiores mares prae caeteris admittunt, ad eos confugiunt, a junioribus agitatae, qui hos in fugam conjiciunt.” (49. ‘Amoenitates Acad.’ vol. iv. 1788, p. 160.) A clergyman, who has bred many pigs, asserts that sows often reject one boar and immediately accept another.

From these facts there can be no doubt that, with most of our domesticated quadrupeds, strong individual antipathies and preferences are frequently exhibited, and much more commonly by the female than by the male. This being the case, it is improbable that the unions of quadrupeds in a state of nature should be left to mere chance. It is much more probable that the females are allured or excited by particular males, who possess certain characters in a higher degree than other males; but what these characters are, we can seldom or never discover with certainty.