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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 (Rhynchæa), 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
species that may be encountered, while they are said even to attack birds of prey that approach too closely to their nests. It must be admitted that these facts agree well with the theory that colour and ornament are due to surplus vital energy and a long course of unchecked development. We have also direct evidence that the males are more active and energetic than the females. Mr. Gosse says that the whirring made by the male Polytmus humming-bird is shriller than that produced by the female ; and he also informs us that the male flies higher and frequents mountains while the female keeps to the lowlands.
Theory of Typical Colours.—The remaining kinds of animal colours, those which can neither be classed as protective, warning, or sexual, are for the most part readily explained on the general principles of the development of colour which we have now laid down. It is a most suggestive fact, that, in cases where colour is required only as a warning, as among the uneatable caterpillars, we find, not one or two glaring tints only, but every kind of colour disposed in elegant patterns, and exhibiting almost as much variety and beauty as among insects and birds. Yet here, not only is sexual selection out of the question, but the need for recognition and identification by others of the same species, seems equally unnecessary. We can then only impute this variety to the normal production of colour in organic forms, when fully exposed to light and air and undergoing great and rapid developmental modification. Among more perfect animals, where the need for recognition has been added, we find intensity and variety of colour at its highest pitch among the South American
butterflies of the families Heliconidæ and Danaidæ, as well as among the Nymphalidæ and Erycinidæ, many of which obtain the necessary protection in other ways. Among birds also, wherever the habits are such that no special protection is needed for the females, and where the species frequent the depths of tropical forests, and are thus naturally protected from the swoop of birds of prey, we find almost equally intense coloration ; as in the trogons, barbets, and gapers.
Local Causes of Colour-development.-Another real, though as yet inexplicable cause of diversity of colour, is to be found in the influence of locality. It is observed that species of totally distinct groups are coloured alike in one district, while in another district the allied species all undergo the same change of colour. Cases of this kind have been adduced by Mr. Bates, by Mr. Darwin, and by myself, and I have collected all the more curious and important examples in my Address to the Biological Section of the British Association, at Glasgow in 1876 (Chap. VII. of this volume). The most probable cause for these simultaneous variations would seem to be the presence of peculiar elements or chemical compounds in the soil, the water, or the atmosphere, or of special organic substances in the vegetation; and a wide field is thus offered for chemical investigation in connection with this interesting subject. Yet, however we may explain it the fact remains, of the same vivid colours in definite patterns being produced in quite unrelated groups, which only agree, so far as we yet know, in inhabiting the same locality.
Summary on Colour-development in Animals.—Let us now sum up the conclusion at which we have arrived,