How Animal Colours are Produced.—The colouring matters of animals are very varied. Copper has been found in the red pigment of the wing of the turaco, and Mr. Sorby has detected no less than seven distinct colouring matters in birds’ eggs, several of which are chemically related to those of blood and bile. The same colours are often produced by quite different substances in different groups, as shown by the red of the wing on the burnet-moth changing to yellow with muriatic acid, while the red of the red-admiral-butterfly undergoes no such change.

These pigmental colours have a different character in animals according to their position in the integument. Following Dr. Hagen’s classification, epidermal colours are those which exist in the external chitinised skin of insects, in the hairs of mammals, and, partially, in the feathers of birds. They are often very deep and rich, and do not fade after death. The hypodermal colours are those which are situated in the inferior soft layer of the skin. These are often of lighter and more vivid tints, and usually fade after death. Many of the reds and yellows of butterflies and birds belong to this class, as well as the intensely vivid hues of the naked skin about the heads of many birds. These colours sometimes exude through the pores, forming an evanescent bloom on the surface.

Interference colours are less frequent in the organic world. They are caused in two ways: either by reflection from the two surfaces of transparent films, as seen in the soap-bubble and in thin films of oil on water; or by fine striæ which produce colours either by reflected or transmitted light, as seen in mother-of-pearl and in finely-ruled metallic surfaces. In both cases colour is produced by light of one wave-length being neutralised, owing to one set of such waves being caused to be half a wave length behind the other set, as may be found explained in any treatise on physical optics. The result is, that the complementary colour of that neutralised is seen; and, as the thickness of the film or the fineness of the striæ undergo slight changes, almost any colour can be produced. This is believed to be the origin of many of the glossy or metallic tints of insects, as well as those of the feathers of some birds. The iridescent colours of the wings of dragon-flies are caused by the superposition of two or more transparent lamellæ; while the shining blue of the Purple-Emperor and other butterflies, and the intensely metallic colours of humming-birds, are probably due to fine striæ.

Colour a Normal Product of Organization.—This outline sketch of the nature of colour in the animal world, however imperfect, will at least serve to show us how numerous and varied are the causes which perpetually tend to the production of colour in animal tissues. If we consider, that in order to produce white, all the rays which fall upon an object must be reflected in the same proportions as they exist in solar light—whereas, if rays of any one or more kinds are absorbed or neutralised, the resultant reflected light will be coloured; and that this colour may be infinitely varied according to the proportions in which different rays are reflected or absorbed—we should expect that white would be, as it really is, comparatively rare and exceptional in nature. The same observation will apply to black, which arises from the absorption of all the different rays. Many of the complex substances which exist in animals and plants are subject to changes of colour under the influence of light, heat, or chemical change, and we know that chemical changes are continually occurring during the physiological processes of development and growth. We also find that every external character is subject to minute changes, which are generally perceptible to us in closely allied species; and we can therefore have no doubt that the extension and thickness of the transparent lamellæ, and the fineness of the striæ or rugosities of the integuments, must be undergoing constant minute changes; and these changes will very frequently produce changes of colour. These considerations render it probable that colour is a normal and even necessary result of the complex structure of animals and plants; and that those parts of an organism which are undergoing continual development and adaptation to new conditions, and are also continually subject to the action of light and heat, will be the parts in which changes of colour will most frequently appear. Now there is little doubt that the external changes of animals and plants in adaptation to the environment are much more numerous than the internal changes; as seen in the varied character of the integuments and appendages of animals—hair, horns, scales, feathers, &c. &c.—and in plants, the leaves, bark, flowers, and fruit, with their various modifications—as compared with the great uniformity in the texture and composition of their internal tissues; and this accords with the uniformity of the tints of blood, muscle, nerve, and bone throughout extensive groups, as compared with the great diversity of colour of their external organs. It seems a fair conclusion that colour per se may be considered to be normal, and to need no special accounting for; while the absence of colour (that is, either white or black), or the prevalence of certain colours to the constant exclusion of others, must be traced, like other modifications in the economy of living things, to the needs of the species. Or, looking at it in another aspect, we may say, that amid the constant variations of animals and plants colour is ever tending to vary and to appear where it is absent; and that natural selection is constantly eliminating such tints as are injurious to the species, or preserving and intensifying such as are useful.

This view is in accordance with the well-known fact, of colours which rarely or never appear in the species in a state of nature, continually occurring among domesticated animals and cultivated plants; showing us that the capacity to develop colour is ever present, so that almost any required tint can be produced which may, under changed conditions, be useful, in however small a degree.

Let us now see how these principles will enable us to understand and explain the varied phenomena of colour in nature, taking them in the order of our functional classification of colours.

Theory of Protective Colours.—We have seen that obscure or protective tints in their infinitely varied degrees are present in every part of the animal kingdom, whole families or genera being often thus coloured. Now the various brown, earthy, ashy, and other neutral tints are those which would be most readily produced, because they are due to an irregular mixture of many kinds of rays; while pure tints require either rays of one kind only, or definite mixtures in proper proportions of two or more kinds of rays. This is well exemplified by the comparative difficulty of producing definite pure tints by the mixture of two or more pigments; while a haphazard mixture of a number of these will be almost sure to produce browns, olives, or other neutral or dingy colours. An indefinite or irregular absorption of some rays and reflection of others would, therefore, produce obscure tints; while pure and vivid colours would require a perfectly definite absorption of one portion of the coloured rays, leaving the remainder to produce the true complementary colour. This being the case we may expect these brown tints to occur when the need of protection is very slight or even when it does not exist at all; always supposing that bright colours are not in any way useful to the species. But whenever a pure colour is protective,—as green in tropical forests or white among arctic snows, there is no difficulty in producing it, by natural selection acting on the innumerable slight variations of tint which are ever occurring. Such variations may, as we have seen, be produced in a great variety of ways; either by chemical changes in the secretions, or by molecular changes in surface structure; and may be brought about by change of food, by the photographic action of light, or by the normal process of generative variation. Protective colours therefore, however curious and complex they may be in certain cases, offer no real difficulties.

Theory of Warning Colours.—These differ greatly from the last class, inasmuch as they present us with a variety of brilliant hues, often of the greatest purity, and combined in striking contrasts and conspicuous patterns. Their use depends upon their boldness and visibility, not on the presence of any one colour; hence we find among these groups some of the most exquisitely-coloured objects in nature. Many of the uneatable caterpillars are strikingly beautiful; while the Danaidæ, Heliconidæ, and protected groups of Papilionidæ, comprise a series of butterflies of the most brilliant and contrasted colours. The bright colours of many of the sea-anemones and sea-slugs will probably be found to be in this sense protective, serving as a warning of their uneatableness. On our theory none of these colours offer any difficulty. Conspicuousness being useful, every variation tending to brighter and purer colours was selected; the result being the beautiful variety and contrast we find.

Imitative Warning Colours:—The Theory of Mimicry.—We now come to those groups which gain protection solely by being mistaken for some of these brilliantly coloured but uneatable creatures, and here a difficulty really exists, and to many minds is so great as to be insuperable. It will be well therefore to endeavour to explain how the resemblance in question may have been brought about.

The most difficult case, and the one which may be taken as a type of the whole class, is that of the genus Leptalis (a group of South American butterflies allied to our common white and yellow kinds), many of the larger species of which are still white or yellow, and which are all eatable by birds and other insectivorous creatures. But there are also a number of species of Leptalis, which are brilliantly red, yellow, and black, and which, band for band and spot for spot, resemble some one of the Danaidæ or Heliconidæ which inhabit the same district and which are nauseous and uneatable. Now the usual difficulty is, that a slight approach to one of these protected butterflies would be of no use, while a greater sudden variation is not admissible on the theory of gradual change by indefinite slight variations. This objection depends almost wholly on the supposition that, when the first steps towards mimicry occurred, the South American Danaidæ were what they are now; while the ancestors of the Leptalides were like the ordinary white or yellow Pieridæ to which they are allied. But the danaioid butterflies of South America are so immensely numerous and so greatly varied, not only in colour but in structure, that we may be sure they are of vast antiquity and have undergone great modification. A large number of them, however, are still of comparatively plain colours, often rendered extremely elegant by the delicate transparency of the wing membrane, but otherwise not at all conspicuous. Many have only dusky or purplish bands or spots; others have patches of reddish or yellowish brown—perhaps the commonest colour among butterflies; while a considerable number are tinged or spotted with yellow, also a very common colour, and one especially characteristic of the Pieridæ, the family to which Leptalis belongs. We may therefore reasonably suppose that in the early stages of the development of the Danaidæ, when they first began to acquire those nauseous secretions which are now their protection, their colours were somewhat plain; either dusky with paler bands and spots, or yellowish with dark borders, and sometimes with reddish bands or spots. At this time they had probably shorter wings and a more rapid flight, just like the other unprotected families of butterflies. But as soon as they became decidedly unpalatable to any of their enemies, it would be an advantage to them to be readily distinguished from all the eatable kinds; and as butterflies were no doubt already very varied in colour, while all probably had wings adapted for rather quick or jerking flight, the best distinction might have been found in outline and habits; whence would arise the preservation of those varieties whose longer wings, bodies, and antennæ, as well as their slower flight, rendered them noticeable—characters which now distinguish the whole group in every part of the world.

Now it would be at this stage, that some of the weaker-flying Pieridæ which happened to resemble some of the Danaidæ around them in their yellow and dusky tints and in the general outline of their wings, would be sometimes mistaken for them by the common enemy, and would thus gain an advantage in the struggle for existence. Admitting this one step to be made, and all the rest must inevitably follow from simple variation and survival of the fittest. So soon as the nauseous butterfly varied in form or colour to such an extent that the corresponding eatable butterfly no longer closely resembled it, the latter would be exposed to attacks, and only those variations would be preserved which kept up the resemblance. At the same time we may well suppose the enemies to become more acute and able to detect smaller differences than at first. This would lead to the destruction of all adverse variations, and thus keep up in continually increasing complexity the outward mimicry which now so amazes us. During the long ages in which this process has been going on, and the Danaidæ have been acquiring those specialities of colour which aid in their preservation, many a Leptalis may have become extinct from not varying sufficiently in the right direction and at the right time to keep up a protective resemblance to its neighbour; and this well accords with the comparatively small number of cases of true mimicry, as compared with the frequency of those protective resemblances to vegetable or inorganic objects whose forms are less definite and colours less changeable. About a dozen other genera of butterflies and moths mimic the Danaidæ in various parts of the world, and exactly the same explanation will apply to all of them. They represent those species of each group which, at the time when the Danaidæ first acquired their protective secretions, happened outwardly to resemble some of them, and which have, by concurrent variation aided by a rigid selection, been able to keep up that resemblance to the present day.[19]

Theory of Sexual Colours.—In Mr. Darwin’s celebrated work, The Descent of Man and Selection in Relation to Sex, he has treated of sexual colour in combination with other sexual characters, and has arrived at the conclusion that all or almost all the colours of the higher animals (including among these insects and all vertebrates) are due to voluntary or conscious sexual selection; and that diversity of colour in the sexes is due, primarily, to the transmission of colour-variations either to one sex only or to both sexes; the difference depending on some unknown law, and not being due to natural selection.

I have long held this portion of Mr. Darwin’s theory to be erroneous; and have argued that the primary cause of sexual diversity of colour was the need of protection, repressing in the female those bright colours which are normally produced in both sexes by general laws; and I have attempted to explain many of the more difficult cases on this principle. (“A Theory of Birds’ Nests,” in Contributions, &c., p. 231.) As I have since given much thought to this subject, and have arrived at some views which appear to me to be of considerable importance, it will be well to sketch briefly the theory I now hold, and afterwards show its application to some of the detailed cases adduced in Mr. Darwin’s work.

The very frequent superiority of the male bird or insect in brightness or intensity of colour, even when the general coloration is the same in both sexes, now seems to me to be, primarily, due to the greater vigour and activity and the higher vitality of the male. The colours of an animal usually fade during disease or weakness, while robust health and vigour adds to their intensity. This is a most important and suggestive fact, and one that appears to hold universally. In all quadrupeds a “dull coat” is indicative of ill-health or low condition; while a glossy coat and sparkling eye are the invariable accompaniments of health and energy. The same rule applies to the feathers of birds, whose colours are only seen in their purity during perfect health; and a similar phenomenon occurs even among insects, for the bright hues of caterpillars begin to fade as soon as they become inactive preparatory to undergoing their transformation. Even in the vegetable kingdom we see the same thing; for the tints of foliage are deepest, and the colours of flowers and fruits richest, on those plants which are in the most healthy and vigorous condition.

This intensity of coloration becomes most developed in the male during the breeding season, when the vitality is at a maximum. It is also very general in those cases in which the male is smaller than the female, as in the hawks and in most butterflies and moths. The same phenomena occur, though in a less marked degree, among mammalia. Whenever there is a difference of colour between the sexes the male is the darker or more strongly marked, and the difference of intensity is most visible during the breeding season (Descent of Man, p. 533). Numerous cases among domestic animals also prove, that there is an inherent tendency in the male to special developments of dermal appendages and colour, quite independently of sexual or any other form of selection. Thus,—“the hump on the male zebu cattle of India, the tail of fat-tailed rams, the arched outline of the forehead in the males of several breeds of sheep, and the mane, the long hairs on the hind legs, and the dew-lap of the male of the Berbura goat,”—are all adduced by Mr. Darwin as instances of characters peculiar to the male, yet not derived from any parent ancestral form. Among domestic pigeons the character of the different breeds is often most strongly manifested in the male birds; the wattles of the carriers and the eye-wattles of the barbs are largest in the males, and male pouters distend their crops to a much greater extent than do the females, while the cock fantails often have a greater number of tail-feathers than the females. There are also some varieties of pigeons of which the males are striped or spotted with black while the females are never so spotted (Animals and Plants under Domestication, I. 161); yet in the parent stock of these pigeons there are no differences between the sexes either of plumage or colour, and artificial selection has not been applied to produce them.

The greater intensity of coloration in the male—which may be termed the normal sexual difference, would be further developed by the combats of the males for the possession of the females. The most vigorous and energetic usually being able to rear most offspring, intensity of colour, if dependent on, or correlated with vigour, would tend to increase. But as differences of colour depend upon minute chemical or structural differences in the organism, increasing vigour acting unequally on different portions of the integument, and often producing at the same time abnormal developments of hair, horns, scales, feathers, &c., would almost necessarily lead also to variable distribution of colour, and thus to the production of new tints and markings. These acquired colours would, as Mr. Darwin has shown, be transmitted to both sexes or to one only, according as they first appeared at an early age, or in adults of one sex; and thus we may account for some of the most marked differences in this respect. With the exception of butterflies, the sexes are almost alike in the great majority of insects. The same is the case in mammals and reptiles; while the chief departure from the rule occurs in birds, though even here in very many cases the law of sexual likeness prevails. But in all cases where the increasing development of colour became disadvantageous to the female, it would be checked by natural selection; and thus produce those numerous instances of protective colouring in the female only, which occur in these two groups, birds and butterflies.

Colour as a Means of Recognition.—There is also, I believe, a very important purpose and use of the varied colours of the higher animals, in the facility it affords for recognition by the sexes or by the young of the same species; and it is this use which probably fixes and determines the coloration in many cases. When differences in size and form are very slight, colour affords the only means of recognition at a distance, or while in motion; and such a distinctive character must therefore be of especial value to flying insects which are continually in motion, and encounter each other, as it were, by accident. This view offers us an explanation of the curious fact, that among butterflies the females of closely-allied species in the same locality sometimes differ considerably, while the males are much alike; for, as the males are the swiftest and by far the highest fliers, and seek out the females, it would evidently be advantageous for them to be able to recognise their true partners at some distance off. This peculiarity occurs with many species of Papilio, Diadema, Adolias, and Colias; and these are all genera, the males of which are strong on the wing and mount high in the air. In birds such marked differences of colour are not required, owing to their higher organization and more perfect senses, which render recognition easy by means of a combination of very slight differential characters.

This principle may perhaps, however, account for some anomalies of coloration among the higher animals. Thus, while admitting that the hare and the rabbit are coloured protectively, Mr. Darwin remarks that the latter while running to its burrow, is made conspicuous to the sportsman, and no doubt to all beasts of prey, by its upturned white tail. But this very conspicuousness while running away, may be useful as a signal and guide to the young, who are thus enabled to escape danger by following the older rabbits, directly and without hesitation, to the safety of the burrow; and this may be the more important from the semi-nocturnal habits of the animal. If this explanation is correct, and it certainly seems probable, it may serve as a warning of how impossible it is, without exact knowledge of the habits of an animal and a full consideration of all the circumstances, to decide that any particular coloration cannot be protective or in any way useful. Mr. Darwin himself is not free from such assumptions. Thus, he says:—“The zebra is conspicuously striped, and stripes cannot afford any protection on the open plains of South Africa.” But the zebra is a very swift animal, and, when in herds, by no means void of means of defence. The stripes therefore may be of use by enabling stragglers to distinguish their fellows at a distance, and they may be even protective when the animal is at rest among herbage—the only time when it would need protective colouring. Until the habits of the zebra have been observed with special reference to these points, it is surely somewhat hasty to declare that the stripes “cannot afford any protection.”

Colour Proportionate to Integumentary Development.—The wonderful display and endless variety of colour in which butterflies and birds so far exceed all other animals, seems primarily due to the excessive development and endless variations of the integumentary structures. No insects have such widely-expanded wings in proportion to their bodies as butterflies and moths; in none do the wings vary so much in size and form, and in none are they clothed with such a beautiful and highly-organized coating of scales. According to the general principles of the production of colour already explained, these long-continued expansions of membranes and developments of surface structures, must have led to numerous colour-changes; which have been sometimes checked, sometimes fixed and utilised, sometimes intensified, by natural selection, according to the needs of the animal. In birds, too, we have the wonderful clothing of plumage—the most highly organized, the most varied, and the most expanded of all dermal appendages. The endless processes of growth and change during the development of feathers, and the enormous extent of this delicately-organized surface, must have been highly favourable to the production of varied colour-effects; which, when not injurious, have been merely fixed for purposes of specific identification, but have often been modified or suppressed whenever different tints were needed for purposes of protection.

Selection by Females not a Cause of Colour.—To conscious sexual selection, that is, the actual choice by the females of the more brilliantly-coloured males, I believe very little if any effect is directly due. It is undoubtedly proved that in birds the females do sometimes exert a choice; but the evidence of this fact collected by Mr. Darwin (Descent of Man, chap. xiv.) does not prove that colour determines that choice, while much of the strongest evidence is directly opposed to this view. All the facts appear to be consistent with the choice depending on a variety of male characteristics, with some of which colour is often correlated. Thus it is the opinion of some of the best observers that vigour and liveliness are most attractive, and these are no doubt usually associated with intensity of colour. Again, the display of the various ornamental appendages of the male during courtship may be attractive; but these appendages, with their bright colours or shaded patterns, are due probably to general laws of growth, and to that superabundant vitality which we have seen to be a cause of colour. But there are many considerations which seem to show that the possession of these ornamental appendages and bright colours in the male is not an important character functionally, and that it has not been produced by the action of conscious sexual selection. Amid the copious mass of facts and opinions collected by Mr. Darwin as to the display of colour and ornaments by the male birds, there is a total absence of any evidence that the females admire or even notice this display. The hen, the turkey, and the pea-fowl go on feeding while the male is displaying his finery; and there is reason to believe that it is his persistency and energy rather than his beauty which wins the day. Again, evidence collected by Mr. Darwin himself proves that each bird finds a mate under any circumstances. He gives a number of cases of one of a pair of birds being shot, and the survivor being always found paired again almost immediately. This is sufficiently explained on the assumption that the destruction of birds by various causes is continually leaving widows and widowers in nearly equal proportions, and thus each one finds a fresh mate; and it leads to the conclusion that permanently unpaired birds are very scarce; so that, speaking broadly, every bird finds a mate and breeds. But this would almost or quite neutralize any effect of sexual selection of colour or ornament, since the less highly-coloured birds would be at no disadvantage as regards leaving healthy offspring. If, however, heightened colour is correlated with health and vigour; and if these healthy and vigorous birds provide best for their young, and leave offspring which, being equally healthy and vigorous, can best provide for themselves—which cannot be denied; then natural selection becomes a preserver and intensifier of colour.

Another most important consideration is, that male butterflies rival or even excel the most gorgeous male birds in bright colours and elegant patterns; and among these there is literally not one particle of evidence that the female is influenced by colour, or even that she has any power of choice; while there is much direct evidence to the contrary (Descent of Man, p. 318). The weakness of the evidence for conscious sexual selection among these insects is so palpable, that Mr. Darwin is obliged to supplement it by the singularly inconclusive argument that, “Unless the female prefer one male to another, the pairing must be left to mere chance, and this does not appear probable” (l.c. p. 317). But he has just said—“The males sometimes fight together in rivalry, and many may be seen pursuing or crowding round the same female;” while in the case of the silk-moths,—“the females appear not to evince the least choice in regard to their partners.” Surely the plain inference from all this is, that males fight and struggle for the almost passive female; and that the most vigorous and energetic, the strongest-winged or the most persevering, wins her. How can there be chance in this? Natural selection would here act, as in birds, in perpetuating the strongest and most vigorous males; and as these would usually be the more highly coloured of their race, the same results would be produced as regards the intensification and variation of colour in the one case as in the other.

Let us now see how these principles will apply to some of the cases adduced by Mr. Darwin in support of his theory of conscious sexual selection.

In Descent of Man, 2nd ed., pp. 307–316, we find an elaborate account of the various modes of colouring of butterflies and moths, proving that the coloured parts are always more or less displayed, and that they have some evident relation to an observer. Mr. Darwin then says: “From the several foregoing facts it is impossible to admit that the brilliant colours of butterflies, and of some few moths, have commonly been acquired for the sake of protection. We have seen that their colours and elegant patterns are arranged and exhibited as if for display. Hence I am led to believe that the females prefer or are most excited by the more brilliant males; for on any other supposition the males would, as far as we can see, be ornamented to no purpose” (l.c., p. 316). I am not aware that any one has ever maintained that the brilliant colours of butterflies have “commonly been acquired for the sake of protection,” yet Mr. Darwin has himself referred to cases in which the brilliant colour is so placed as to serve for protection; as for example, the eye-spots on the hind wings of moths, which are pierced by birds and so save the vital parts of the insect; while the bright patch on the orange-tip butterflies which Mr. Darwin denies are protective, may serve the same purpose. It is in fact somewhat remarkable how very generally the black spots, ocelli, or bright patches of colour are on the tips, margins, or discs of the wings; and as the insects are necessarily visible while flying, and this is the time when they are most subject to attacks by insectivorous birds, the position of the more conspicuous parts at some distance from the body may be a real protection to them. Again, Mr. Darwin admits that the white colour of the male ghost-moth may render it more easily seen by the female while flying about in the dusk; and if to this we add that it will be also more readily distinguished, from allied species, we have a reason for diverse ornamentation in these insects quite sufficient to account for most of the facts, without believing in the selection of brilliant males by the females, for which there is not a particle of evidence.[20]

Probable use of the Horns of Beetles.—A somewhat analogous case is furnished by the immense horns of some beetles of the families Copridæ and Dynastidæ, which Mr. Darwin admits are not used for fighting, and therefore concludes are ornaments, developed through selection of the larger-horned males by the females. But it has been overlooked that these horns may be protective. The males probably fly about most, as is usually the case with male insects; and as they generally fly at dusk they are subject to the attacks of large-mouthed goatsuckers and podargi, as well as of insect-eating owls. Now the long, pointed or forked horns, often divergent, or movable with the head, would render it very difficult for these birds to swallow such insects, and would therefore be an efficient protection; just as are the hooked spines of some stingless ants and the excessively hard integuments of many beetles, against the smaller insectivorous birds.

Cause of the greater Brilliancy of some Female Insects.—The facts given by Mr. Darwin to show that butterflies and other insects can distinguish colours and are attracted by colours similar to their own, are quite consistent with the view that colour, which continually tends to appear, is utilised for purposes of identification and distinction, when not required to be modified or suppressed for the purpose of protection. The cases of the females of some species of Thecla, Callidryas, Colias, and Hipparchia, which have more conspicuous markings than the male, may be due to several causes: to obtain greater distinction from other species; for protection from birds, as in the case of the yellow-underwing moths; while sometimes—as in Hipparchia—the lower intensity of colouring in the female may lead to more contrasted markings. Mr. Darwin thinks that here the males have selected the more beautiful females; although one chief fact in support of his theory of conscious sexual selection is, that throughout the whole animal kingdom the males are usually so ardent that they will accept any female, while the females are coy, and choose the handsomest males, whence it is believed the general brilliancy of males as compared with females has arisen.

Perhaps the most curious cases of sexual difference of colour are those in which the female is very much more gaily coloured than the male. This occurs most strikingly in some species of Pieris in South America, and of Diadema in the Malay islands; and in both cases the females resemble species of the uneatable Danaidæ and Heliconidæ, and thus gain a protection. In the case of Pieris pyrrha, P. malenka, and P. lorena, the males are plain white and black, while the females are orange, yellow, and black, and so banded and spotted as exactly to resemble species of Heliconidæ. Mr. Darwin admits that these bright colours have been acquired for protection; but as there is no apparent cause for the strict limitation of the colour to the female, he believes that it has been kept down in the male by its being unattractive to her. This appears to me to be a supposition opposed to the whole theory of sexual selection itself. For this theory is, that minute variations of colour in the male are attractive to the female, have always been selected, and that thus the brilliant male colours have been produced. But in this case he thinks that the female butterfly had a constant aversion to every trace of colour, even when we must suppose it was constantly recurring during the successive variations which resulted in such a marvellous change in herself. But the case admits of a much more simple interpretation. For if we consider the fact that the females frequent the forests where the Heliconidæ abound, while the males fly much in the open and assemble in great numbers with other white and yellow butterflies on the banks of rivers; may it not be possible that the appearance of orange stripes or patches would be as injurious to the male as it is useful to the female, by making him a more easy mark for insectivorous birds among his white companions? This seems a more probable supposition, than the altogether hypothetical choice of the female, sometimes exercised in favour of and sometimes against every new variety of colour in her partner.

A strictly analogous case is that of the glow-worm, whose light, as originally suggested by Mr. Belt, is admitted to be a warning of its uneatability to insectivorous nocturnal animals. The male, having wings, does not require this protection. In the tropics the number of nocturnal insectivorous birds and bats is very much greater, hence winged species possess the light, as they would otherwise be eaten by mistake for more savoury insects; and it may be that the luminous Elateridæ of the tropics really mimic the true fireflies (Lampyridæ), which are uneatable. This is the more probable as the Elateridæ, in the great majority of species, have brown or protective colours, and are therefore certainly palatable to insectivorous animals.

Origin of the Ornamental Plumage of Male Birds.—We now come to such wonderful developments of plumage and colour as are exhibited by the peacock and the Argus-pheasant; and I may here mention that it was the case of the latter bird, as fully discussed by Mr. Darwin, which first shook my belief in “sexual,” or more properly “female” selection. The long series of gradations, by which the beautifully shaded ocelli on the secondary wing-feathers of this bird, have been produced, are clearly traced out; the result being a set of markings, so exquisitely shaded as to represent “balls lying loose within sockets”—purely artificial objects of which these birds could have no possible experience. That this result should have been attained through thousands and tens of thousands of female birds all preferring those males whose markings varied slightly in this one direction, this uniformity of choice continuing through thousands and tens of thousands of generations, is to me absolutely incredible. And when, further, we remember that those which did not so vary, would also, according to all the evidence, find mates and leave offspring, the actual result seems quite impossible of attainment by such means.

Without pretending to solve completely so difficult a problem as that of the origin and uses of the variously coloured plumes and ornaments so often possessed by male birds, I would point out a few facts which seem to afford a clue. And first, the most highly-coloured and most richly-varied markings occur on those parts of the plumage which have undergone the greatest modification, or have acquired the most abnormal development. In the peacock, the tail-coverts are enormously developed, and the “eyes” are situated on the greatly dilated ends. In the birds-of-paradise, breast, or neck, or head, or tail-feathers, are greatly developed and highly coloured. The hackles of the cock, and the scaly breasts of humming-birds are similar developments; while in the Argus-pheasant the secondary quills are so enormously lengthened and broadened as to have become almost useless for flight. Now it is easily conceivable, that during this process of development, inequalities in the distribution of colour may have arisen in different parts of the same feather; and that spots and bands may thus have become broadened out into shaded spots or ocelli, in the way indicated by Mr. Darwin, much as the spots and rings on a soap-bubble increase with increasing tenuity. This is the more probable, because in domestic fowls varieties of colour tend to become symmetrical, quite independently of sexual selection. (Descent of Man, p. 424.)

If now we accept the evidence of Mr. Darwin’s most trustworthy correspondents, that the choice of the female, so far as she exerts any, falls upon the “most vigorous, defiant, and mettlesome male;” and if we further believe, what is certainly the case, that these are as a rule the most brightly coloured and adorned with the finest developments of plumage, we have a real and not a hypothetical cause at work. For these most healthy, vigorous, and beautiful males will have the choice of the finest and most healthy females; will have the most numerous and healthy families; and will be able best to protect and rear those families. Natural selection, and what may be termed male selection, will tend to give them the advantage in the struggle for existence; and thus the fullest plumage and the finest colours will be transmitted, and tend to advance in each succeeding generation.

Theory of Display of Ornaments by Males.—The full and interesting account given by Mr. Darwin of the colours and habits of male and female birds (Descent of Man, Chapters xiii. and xiv.), proves that in most, if not in all cases, the male birds fully display their ornamental plumage before the females or in rivalry with each other; but on the essential point of whether the female’s choice is determined by minute differences in these ornaments or in their colours, there appears to be an entire absence of evidence. In the section on “Preference for particular Males by the Females,” the facts quoted show indifference to colour, except that some colour similar to their own seems to be preferred. But in the case of the hen canary, who chose a greenfinch in preference to either chaffinch or goldfinch, gay colours had evidently no preponderating attraction. There is some evidence adduced that female birds may, and probably do, choose their mates; but none whatever that the choice is determined by difference of colour; and no less than three eminent breeders informed Mr. Darwin that they “did not believe that the females prefer certain males on account of the beauty of their plumage.” Again, Mr. Darwin himself says: “As a general rule colour appears to have little influence on the pairing of pigeons.” The oft-quoted case of Sir R. Heron’s pea-hens which preferred an “old pied cock” to those normally coloured, is a very unfortunate one; because pied birds are just those that are not favoured in a state of nature, or the breeds of wild animals would become as varied and mottled as our domestic varieties. If such irregular fancies were not rare exceptions, the production of definite colours and patterns by the choice of the female birds, or in any other way, would be impossible.

There remains, however, to be accounted for, the remarkable fact of the display by the male of each species of its peculiar beauties of plumage and colour,—a display which Mr. Darwin evidently considers his strongest argument in favour of conscious selection by the female. This display is, no doubt, a very interesting and important phenomenon; but it may, I believe, be satisfactorily explained on the general principles here laid down, without calling to our aid a purely hypothetical choice exerted by the female bird.

At pairing-time, the male is in a state of excitement, and full of exuberant energy. Even unornamental birds flutter their wings or spread them out, erect their tails or crests, and thus give vent to the nervous excitability with which they are overcharged. It is not improbable that crests and other erectile feathers may be primarily of use in frightening away enemies, since they are generally erected when angry or during combat. Those individuals who were most pugnacious and defiant, and who brought these erectile plumes most frequently and most powerfully into action, would tend to increase them by use, and to leave them further developed in some of their descendants. If, in the course of this development, colour appeared—and we have already shown that such developments of plumage are a very probable cause of colour—we have every reason to believe it would be most vivid in these most pugnacious and energetic individuals; and as these would always have the advantage in the rivalry for mates (to which advantage the excess of colour and plumage might sometimes conduce), there seems nothing to prevent a progressive development of these ornaments in all dominant races; that is, wherever there was such a surplus of vitality, and such 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, five-sixths, 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 (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 some 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 beautifully-coloured 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, as to the various modes in which colour is produced or modified in the animal kingdom.

The various causes of colour in the animal world are, molecular and chemical change of the substance of their integuments, or the action on it of heat, light or moisture. It is also produced by interference of light in superposed transparent lamellæ, or by excessively fine surface-striæ. These elementary conditions for the production of colour are found everywhere in the surface-structures of animals, so that its presence must be looked upon as normal, its absence as exceptional.

Colours are fixed or modified in animals by natural selection for various purposes; obscure or imitative colours for concealment; gaudy colours as a warning; and special markings, either for easy recognition by strayed individuals, females, or young, or to direct attack from a vital part, as in the large brilliantly-marked wings of some butterflies and moths.

Colours are produced or intensified by processes of development,—either where the integument or its appendages undergo great extension or modification, or where there is a surplus of vital energy, as in male animals generally, and more especially at the breeding season.

Colours are also more or less influenced by a variety of causes, such as the nature of the food, the photographic action of light, and also by some unknown local action probably dependent on chemical peculiarities in the soil or vegetation.

These various causes have acted and reacted in a variety of ways, and have been modified by conditions dependent on age or on sex, on competition with new forms, or on geographical or climatic changes. In so complex a subject, for which experiment and systematic inquiry has done so little, we cannot expect to explain every individual case, or solve every difficulty; but it is believed that all the great features of animal coloration and many of the details become explicable on the principles we have endeavoured to lay down.

It will perhaps be considered presumptuous to put forth this sketch of the subject of colour in animals, as a substitute for one of Mr. Darwin’s most highly elaborated theories—that of voluntary or perceptive sexual selection; yet I venture to think that it is more in accordance with the whole of the facts, and with the theory of natural selection itself; and I would ask such of my readers as may be sufficiently interested in the subject, to read again Chapters XI. to XVI. of the Descent of Man, and consider the whole subject from the point of view here laid down. The explanation of almost all the ornaments and colours of birds and insects as having been produced by the perceptions and choice of the females, has, I believe, staggered many evolutionists, but has been provisionally accepted because it was the only theory that even attempted to explain the facts. It may perhaps be a relief to some of them, as it has been to myself, to find that the phenomena can be shown to depend on the general laws of development, and on the action of “natural selection,” which theory will, I venture to think, be relieved from an abnormal excrescence and gain additional vitality, by the adoption of the views here imperfectly set forth.

Although we have arrived at the conclusion that tropical light and heat can in no sense be considered as the cause of colour, there remains to be explained the undoubted fact that all the more intense and gorgeous tints are manifested by the animal life of the tropics; while in some groups, such as butterflies and birds, there is a marked preponderance of highly-coloured species. This is probably due to a variety of causes, some of which we can indicate, while others remain to be discovered. The luxuriant vegetation of the tropics throughout the entire year affords so much concealment, that colour may there be safely developed to a much greater extent than in climates where the trees are bare in winter, during which season the struggle for existence is most severe, and even the slightest disadvantage may prove fatal. Equally important, probably, has been the permanence of favourable conditions in the tropics, allowing certain groups to continue dominant for long periods, and thus to carry out in one unbroken line whatever developments of plumage or colour may once have acquired an ascendency. Changes of climatal conditions, and pre-eminently the glacial epoch, probably led to the extinction of a host of highly-developed and finely-coloured insects and birds in temperate zones; just as we know that it led to the extinction of the larger and more powerful mammalia which formerly characterised the temperate zone in both hemispheres; and this view is supported by the fact that it is amongst those groups only which are now exclusively tropical that all the more extraordinary developments of ornament and colour are found. The obscure local causes of colour to which we have referred will also have acted most efficiently in regions where the climatal condition remained constant, and where migration was unnecessary; while whatever direct effect may be produced by light or heat, will necessarily have acted more powerfully within the tropics. And lastly, all these causes have been in action over an actually greater area in tropical than in temperate zones; while estimated potentially, in proportion to its life-sustaining power, the lands which enjoy a practically tropical climate (extending as they do considerably beyond the geographical tropics) are very much larger than the temperate regions of the earth.

Combining the effects of all these various causes we are quite able to understand the superiority of the tropical parts of the globe, not only in the abundance and variety of their forms of life, but also as regards the ornamental appendages and vivid coloration which these forms present.