Kitabı oku: «The Descent of Man, and Selection in Relation to Sex», sayfa 41

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.

CHAPTER 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.