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complete adaption to conditions, that the inconvenience or danger produced by such ornaments was so comparatively small as not to affect the superiority of the race over its nearest allies.

But if those portions of the plumage, which were originally erected under the influence of anger or fear, became largely developed and brightly coloured, the actual display, under the influence of jealousy or sexual excitement becomes quite intelligible. The males, in their rivalry with each other, would see what plumes were most effective; and each would endeavour to excel his enemy as far as voluntary exertion would enable him, just as they endeavour to rival each other in song, even sometimes to the point of causing their own destruction.

Natural Selection as Neutralizing Sexual Selection. -There is also a general argument against Mr. Darwin's views on this question, founded on the nature and potency of "natural" as opposed to "sexual" selection, which appears to me to be of itself almost conclusive as to the whole matter at issue. Natural selection, or the survival of the fittest, acts perpetually and on an enormous scale. Taking the offspring of each pair of birds as, on the average, only six annually, one-third of these at most will be preserved, while the two-thirds which are least fitted will die. At intervals of a few years, whenever unfavourable conditions occur, fivesixths, nine-tenths, or even a greater proportion of the whole yearly production are weeded out, leaving only the most perfect and best adapted to survive. Now unless these survivors are, on the whole, the most ornamental, this rigid natural selection must neutralise and destroy any influence that may be exerted by

female selection.

The utmost that can be claimed for

the latter is, that a small fraction of the least ornamented do not obtain mates, while a few of the most ornamented may leave more than the average number of offspring. Unless, therefore, there is the strictest correlation between ornament and general perfection, the more brightly coloured or ornamented varieties can obtain no permanent advantage; and if there is (as I maintain) such a correlation, then the sexual selection. of colour or ornament, for which there is little or no evidence, becomes needless, because natural selection which is an admitted vera causa, will itself produce all the results.

In the case of butterflies the argument becomes even stronger, because the fertility is so much greater than in birds, and the weeding-out of the unfit takes place, to a great extent, in the egg and larva state. Unless the eggs and larvæ which escaped to produce the next generation were those which would produce the more highly-coloured butterflies, it is difficult to perceive how the slight preponderance of colour sometimes selected by the females, should not be wholly neutralized by the extremely rigid selection for other qualities to which the offspring in every stage are exposed. The only way in which we can account for the observed facts is, by the supposition that colour and ornament are strictly correlated with health, vigour, and general fitness to survive. We have shown that there is reason to believe that this is the case, and if so, conscious sexual selection becomes as unnecessary as it would certainly be ineffective.

Greater Brilliancy of some Female Birds.-There is one other very curious case of sexual colouring among

birds-that, namely, in which the female is decidedly brighter or more strongly marked than the male; as in the fighting quails (Turnix), painted snipe (Rhynchoa), two species of phalarope (Phalaropus), and the common cassowary (Casuarius galeatus). In all these cases, it is known that the males take charge of and incubate the eggs, while the females are almost always larger and more pugnacious.

In my "Theory of Birds' Nests" (Natural Selection, p. 251), I imputed this difference of colour to the greater need for protection by the male bird while incubating; to which Mr. Darwin has objected that the difference is not sufficient, and is not always so distributed as to be most effective for this purpose; and he believes that it is due to reversed sexual selection, that is, to the female taking the usual rôle of the male, and being chosen for her brighter tints. We have already seen reason for rejecting this latter theory in every case; and I also admit that Mr. Darwin's criticism is sound, and that my theory of protection is, in this case, only partially, if at all, applicable. But the theory now advanced, of intensity of colour being due to general vital energy, is quite applicable; and the fact that the superiority of the female in this respect is quite exceptional, and is therefore probably not in any case of very ancient date, will account for the difference of colour thus produced being always very slight.

Colour-development as Illustrated by Humming-birds. -Of the mode of action of the general principles of colour-development among animals, we have an excellent example in the humming-birds. Of all birds these are at once the smallest, the most active, and the fullest of

vital energy. When poised in the air their wings are invisible, owing to the rapidity of their motion, and when startled they dart away with the rapidity of a flash of light. Such active creatures would not be an easy prey to any rapacious bird; and if one at length was captured, the morsel obtained would hardly repay the labour. We may be sure, therefore, that they are practically unmolested. The immense variety they exhibit in structure, plumage, and colour, indicates a high antiquity for the race; while their general abundance in individuals shows that they are a dominant group, well adapted to all the conditions of their existence. Here we find everything necessary for the development of colour and accessory plumes. The surplus vital energy shown in their combats and excessive activity, has expended itself in ever-increasing developments of plumage, and greater and greater intensity of colour, regulated only by the need for specific identification which would be especially required in such small and mobile creatures. Thus may be explained those remarkable differences of colour between closely-allied species, one having a crest like the topaz, while in another it resembles the sapphire. The more vivid colours and more developed plumage of the males, I am now inclined to think may be wholly due to their greater vital energy, and to those general laws which lead to such superior developments even in domestic breeds; but in some cases the need of protection by the female while incubating, to which I formerly imputed the whole phenomenon, may have suppressed a portion of the ornament which she would otherwise have attained.

The extreme pugnacity of humming-birds has been

noticed by all observers, and it seems to be to come extent proportioned to the degree of colour and ornament in the species. Thus Mr. Salvin observes of Eugenes fulgens, that it is "a most pugnacious bird," and that 'hardly any species shows itself more brilliantly on the wing." Again of Campylopterus hemileucurus," the pugnacity of this species is remarkable. It is very seldom that two males meet without an aërial battle," and "the large and showy tail of this humming-bird makes it one of the most conspicuous on the wing." Again, the elegant frill-necked Lophornis ornatus "is very pugnacious, erecting its crest, throwing out its whiskers and attacking every humming-bird that may pass within its range of vision;" and of another species L. magnificus, it is said that "it is so bold that the sight of man creates no alarm." The beautifullycoloured Thaumastura Cora "rarely permits any other humming-bird to remain in its neighbourhood, but wages a continual and terrible war upon them." The magnificent bar-tail, Cometes sparganurus, one of the most imposing of all the humming-birds, is extremely fierce and pugnacious, "the males chasing each other through the air with surprising perseverance and acrimony." These are all the species I find noticed as being especially pugnacious, and every one of them is exceptionally coloured or ornamented; while not one of the small, plain, and less ornamental species are so described, although many of them are common and well observed species. It is also to be noticed that the remarkable pugnacity of these birds is not confined to one season or even to birds of the same species, as is usual in sexual combats, but extends to any other

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