The Descent of Man – Day 105 of 151

Variability of Birds, and Especially of Their Secondary Sexual Characters

Variability and inheritance are the foundations for the work of selection. That domesticated birds have varied greatly, their variations being inherited, is certain. That birds in a state of nature have been modified into distinct races is now universally admitted. (33. According to Dr. Blasius (‘Ibis,’ vol. ii. 1860, p. 297), there are 425 indubitable species of birds which breed in Europe, besides sixty forms, which are frequently regarded as distinct species. Of the latter, Blasius thinks that only ten are really doubtful, and that the other fifty ought to be united with their nearest allies; but this shews that there must be a considerable amount of variation with some of our European birds. It is also an unsettled point with naturalists, whether several North American birds ought to be ranked as specifically distinct from the corresponding European species. So again many North American forms which until lately were named as distinct species, are now considered to be local races.) Variations may be divided into two classes; those which appear to our ignorance to arise spontaneously, and those which are directly related to the surrounding conditions, so that all or nearly all the individuals of the same species are similarly modified. Cases of the latter kind have recently been observed with care by Mr. J.A. Allen (34. ‘Mammals and Birds of East Florida,’ also an ‘Ornithological Reconnaissance of Kansas,’ etc. Notwithstanding the influence of climate on the colours of birds, it is difficult to account for the dull or dark tints of almost all the species inhabiting certain countries, for instance, the Galapagos Islands under the equator, the wide temperate plains of Patagonia, and, as it appears, Egypt (see Mr. Hartshorne in the ‘American Naturalist,’ 1873, p. 747). These countries are open, and afford little shelter to birds; but it seems doubtful whether the absence of brightly coloured species can be explained on the principle of protection, for on the Pampas, which are equally open, though covered by green grass, and where the birds would be equally exposed to danger, many brilliant and conspicuously coloured species are common. I have sometimes speculated whether the prevailing dull tints of the scenery in the above named countries may not have affected the appreciation of bright colours by the birds inhabiting them.), who shews that in the United States many species of birds gradually become more strongly coloured in proceeding southward, and more lightly coloured in proceeding westward to the arid plains of the interior. Both sexes seem generally to be affected in a like manner, but sometimes one sex more than the other. This result is not incompatible with the belief that the colours of birds are mainly due to the accumulation of successive variations through sexual selection; for even after the sexes have been greatly differentiated, climate might produce an equal effect on both sexes, or a greater effect on one sex than on the other, owing to some constitutional difference.

Individual differences between the members of the same species are admitted by every one to occur under a state of nature. Sudden and strongly marked variations are rare; it is also doubtful whether if beneficial they would often be preserved through selection and transmitted to succeeding generations. (35. ‘Origin of Species’ fifth edit. 1869, p.104. I had always perceived, that rare and strongly-marked deviations of structure, deserving to be called monstrosities, could seldom be preserved through natural selection, and that the preservation of even highly-beneficial variations would depend to a certain extent on chance. I had also fully appreciated the importance of mere individual differences, and this led me to insist so strongly on the importance of that unconscious form of selection by man, which follows from the preservation of the most valued individuals of each breed, without any intention on his part to modify the characters of the breed. But until I read an able article in the ‘North British Review’ (March 1867, p. 289, et seq.), which has been of more use to me than any other Review, I did not see how great the chances were against the preservation of variations, whether slight or strongly pronounced, occurring only in single individuals.) Nevertheless, it may be worth while to give the few cases which I have been able to collect, relating chiefly to colour,–simple albinism and melanism being excluded. Mr. Gould is well known to admit the existence of few varieties, for he esteems very slight differences as specific; yet he states (36. ‘Introduction to the Trochlidae,’ p. 102.) that near Bogota certain humming-birds belonging to the genus Cynanthus are divided into two or three races or varieties, which differ from each other in the colouring of the tail–“some having the whole of the feathers blue, while others have the eight central ones tipped with beautiful green.” It does not appear that intermediate gradations have been observed in this or the following cases. In the males alone of one of the Australian parrakeets “the thighs in some are scarlet, in others grass-green.” In another parrakeet of the same country “some individuals have the band across the wing-coverts bright-yellow, while in others the same part is tinged with red. (37. Gould, ‘Handbook to Birds of Australia,’ vol. ii. pp. 32 and 68.) In the United States some few of the males of the scarlet tanager (Tanagra rubra) have “a beautiful transverse band of glowing red on the smaller wing-coverts” (38. Audubon, ‘Ornithological Biography,’ 1838, vol. iv. p. 389.); but this variation seems to be somewhat rare, so that its preservation through sexual selection would follow only under usually favourable circumstances. In Bengal the Honey buzzard (Pernis cristata) has either a small rudimental crest on its head, or none at all: so slight a difference, however, would not have been worth notice, had not this same species possessed in Southern India a well-marked occipital crest formed of several graduated feathers.” (39. Jerdon, ‘Birds of India,’ vol. i. p. 108; and Mr. Blyth, in ‘Land and Water,’ 1868, p. 381.)

The following case is in some respects more interesting. A pied variety of the raven, with the head, breast, abdomen, and parts of the wings and tail-feathers white, is confined to the Feroe Islands. It is not very rare there, for Graba saw during his visit from eight to ten living specimens. Although the characters of this variety are not quite constant, yet it has been named by several distinguished ornithologists as a distinct species. The fact of the pied birds being pursued and persecuted with much clamour by the other ravens of the island was the chief cause which led Brunnich to conclude that they were specifically distinct; but this is now known to be an error. (40. Graba, ‘Tagebuch Reise nach Faro,’ 1830, ss. 51-54. Macgillivray, ‘History of British Birds,’ vol. iii. p. 745, ‘Ibis,’ vol. v. 1863, p. 469.) This case seems analogous to that lately given of albino birds not pairing from being rejected by their comrades.

In various parts of the northern seas a remarkable variety of the common Guillemot (Uria troile) is found; and in Feroe, one out of every five birds, according to Graba’s estimation, presents this variation. It is characterised (41. Graba, ibid. s. 54. Macgillivray, ibid. vol. v. p. 327.) by a pure white ring round the eye, with a curved narrow white line, an inch and a half in length, extending back from the ring. This conspicuous character has caused the bird to be ranked by several ornithologists as a distinct species under the name of U. lacrymans, but it is now known to be merely a variety. It often pairs with the common kind, yet intermediate gradations have never been seen; nor is this surprising, for variations which appear suddenly, are often, as I have elsewhere shewn (42. ‘Variation of Animals and Plants under Domestication,’ vol. ii. p. 92.), transmitted either unaltered or not at all. We thus see that two distinct forms of the same species may co-exist in the same district, and we cannot doubt that if the one had possessed any advantage over the other, it would soon have been multiplied to the exclusion of the latter. If, for instance, the male pied ravens, instead of being persecuted by their comrades, had been highly attractive (like the above pied peacock) to the black female ravens their numbers would have rapidly increased. And this would have been a case of sexual selection.

With respect to the slight individual differences which are common, in a greater or less degree, to all the members of the same species, we have every reason to believe that they are by far the most important for the work of selection. Secondary sexual characters are eminently liable to vary, both with animals in a state of nature and under domestication. (43. On these points see also ‘Variation of Animals and Plants under Domestication,’ vol. i. p. 253; vol ii. pp. 73, 75.) There is also reason to believe, as we have seen in our eighth chapter, that variations are more apt to occur in the male than in the female sex. All these contingencies are highly favourable for sexual selection. Whether characters thus acquired are transmitted to one sex or to both sexes, depends, as we shall see in the following chapter, on the form of inheritance which prevails.

It is sometimes difficult to form an opinion whether certain slight differences between the sexes of birds are simply the result of variability with sexually-limited inheritance, without the aid of sexual selection, or whether they have been augmented through this latter process. I do not here refer to the many instances where the male displays splendid colours or other ornaments, of which the female partakes to a slight degree; for these are almost certainly due to characters primarily acquired by the male having been more or less transferred to the female. But what are we to conclude with respect to certain birds in which, for instance, the eyes differ slightly in colour in the two sexes? (44. See, for instance, on the irides of a Podica and Gallicrex in ‘Ibis,’ vol. ii. 1860, p. 206; and vol. v. 1863, p. 426.) In some cases the eyes differ conspicuously; thus with the storks of the genus Xenorhynchus, those of the male are blackish-hazel, whilst those of the females are gamboge-yellow; with many hornbills (Buceros), as I hear from Mr. Blyth (45. See also Jerdon, ‘Birds of India,’ vol. i. pp. 243-245.), the males have intense crimson eyes, and those of the females are white. In the Buceros bicornis, the hind margin of the casque and a stripe on the crest of the beak are black in the male, but not so in the female. Are we to suppose that these black marks and the crimson colour of the eyes have been preserved or augmented through sexual selection in the males? This is very doubtful; for Mr. Bartlett shewed me in the Zoological Gardens that the inside of the mouth of this Buceros is black in the male and flesh-coloured in the female; and their external appearance or beauty would not be thus affected. I observed in Chile (46. ‘Zoology of the Voyage of H.M.S. “Beagle,”‘ 1841, p. 6.) that the iris in the condor, when about a year old, is dark-brown, but changes at maturity into yellowish-brown in the male, and into bright red in the female. The male has also a small, longitudinal, leaden-coloured, fleshy crest or comb. The comb of many gallinaceous birds is highly ornamental, and assumes vivid colours during the act of courtship; but what are we to think of the dull-coloured comb of the condor, which does not appear to us in the least ornamental? The same question may be asked in regard to various other characters, such as the knob on the base of the beak of the Chinese goose (Anser cygnoides), which is much larger in the male than in the female. No certain answer can be given to these questions; but we ought to be cautious in assuming that knobs and various fleshy appendages cannot be attractive to the female, when we remember that with savage races of man various hideous deformities–deep scars on the face with the flesh raised into protuberances, the septum of the nose pierced by sticks or bones, holes in the ears and lips stretched widely open–are all admired as ornamental.

Whether or not unimportant differences between the sexes, such as those just specified, have been preserved through sexual selection, these differences, as well as all others, must primarily depend on the laws of variation. On the principle of correlated development, the plumage often varies on different parts of the body, or over the whole body, in the same manner. We see this well illustrated in certain breeds of the fowl. In all the breeds the feathers on the neck and loins of the males are elongated, and are called hackles; now when both sexes acquire a top-knot, which is a new character in the genus, the feathers on the head of the male become hackle-shaped, evidently on the principle of correlation; whilst those on the head of the female are of the ordinary shape. The colour also of the hackles forming the top-knot of the male, is often correlated with that of the hackles on the neck and loins, as may be seen by comparing these feathers in the golden and silver-spangled Polish, the Houdans, and Creve-coeur breeds. In some natural species we may observe exactly the same correlation in the colours of these same feathers, as in the males of the splendid Gold and Amherst pheasants.

The structure of each individual feather generally causes any change in its colouring to be symmetrical; we see this in the various laced, spangled, and pencilled breeds of the fowl; and on the principle of correlation the feathers over the whole body are often coloured in the same manner. We are thus enabled without much trouble to rear breeds with their plumage marked almost as symmetrically as in natural species. In laced and spangled fowls the coloured margins of the feathers are abruptly defined; but in a mongrel raised by me from a black Spanish cock glossed with green, and a white game-hen, all the feathers were greenish-black, excepting towards their extremities, which were yellowish-white; but between the white extremities and the black bases, there was on each feather a symmetrical, curved zone of dark-brown. In some instances the shaft of the feather determines the distribution of the tints; thus with the body-feathers of a mongrel from the same black Spanish cock and a silver-spangled Polish hen, the shaft, together with a narrow space on each side, was greenish-black, and this was surrounded by a regular zone of dark-brown, edged with brownish-white. In these cases we have feathers symmetrically shaded, like those which give so much elegance to the plumage of many natural species. I have also noticed a variety of the common pigeon with the wing-bars symmetrically zoned with three bright shades, instead of being simply black on a slaty-blue ground, as in the parent-species.

In many groups of birds the plumage is differently coloured in the several species, yet certain spots, marks, or stripes are retained by all. Analogous cases occur with the breeds of the pigeon, which usually retain the two wing-bars, though they may be coloured red, yellow, white, black, or blue, the rest of the plumage being of some wholly different tint. Here is a more curious case, in which certain marks are retained, though coloured in a manner almost exactly the opposite of what is natural; the aboriginal pigeon has a blue tail, with the terminal halves of the outer webs of the two outer tail feathers white; now there is a sub-variety having a white instead of a blue tail, with precisely that part black which is white in the parent-species. (47. Bechstein, ‘Naturgeschichte Deutschlands,’ B. iv. 1795, s. 31, on a sub-variety of the Monck pigeon.)

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