Darwinism (1889) Part 22

During excitement, and when the organism develops superabundant energy, many animals find it pleasurable to exercise their various muscles, often in fantastic ways, as seen in the gambols of kittens, lambs, and other young animals. But at the time of pairing, male birds are in a state of the most perfect development, and possess an enormous store of vitality; and under the excitement of the s.e.xual pa.s.sion they perform strange antics or rapid flights, as much probably from an internal impulse to motion and exertion as with any desire to please their mates.

Such are the rapid descent of the snipe, the soaring and singing of the lark, and the dances of the c.o.c.k-of-the-rock and of many other birds.

It is very suggestive that similar strange movements are performed by many birds which have no ornamental plumage to display. Goatsuckers, geese, carrion vultures, and many other birds of plain plumage have been observed to dance, spread their wings or tails, and perform strange love-antics. The courtship of the great albatross, a most unwieldy and dull coloured bird, has been thus described by Professor Moseley: "The male, standing by the female on the nest, raises his wings, spreads his tail and elevates it, throws up his head with the bill in the air, or stretches it straight out, or forwards, as far as he can, and then utters a curious cry."[127] Mr. Jenner Weir informs me that "the male blackbird is full of action, spreads out his glossy wing and tail, turns his rich golden beak towards the female, and chuckles with delight,"

while he has never seen the more plain coloured thrush demonstrative to the female. The linnet distends his rosy breast, and slightly expands his brown wings and tail; while the various gay coloured Australian finches adopt such att.i.tudes and postures as, in every case, to show off their variously coloured plumage to the best advantage.[128]

_A Theory of Animal Coloration._

Having rejected Mr. Darwin's theory of female choice as incompetent to account for the brilliant colours and markings of the higher animals, the preponderance of these colours and markings in the male s.e.x, and their display during periods of activity or excitement, I may be asked what explanation I have to offer as a preferable subst.i.tute. In my _Tropical Nature_ I have already indicated such a theory, which I will now briefly explain, supporting it by some additional facts and arguments, which appear to me to have great weight, and for which I am mainly indebted to a most interesting and suggestive posthumous work by Mr. Alfred Tylor.[129]

The fundamental or ground colours of animals ar has been shown in preceding chapters, very largely protective, and it is not improbable that the primitive colours of all animals were so. During the long course of animal development other modes of protection than concealment by harmony of colour arose, and thenceforth the normal development of colour due to the complex chemical and structural changes ever going on in the organism, had full play; and the colours thus produced were again and again modified by natural selection for purposes of warning, recognition, mimicry, or special protection, as has been already fully explained in the preceding chapters.

Mr. Taylor has, however, called attention to an important principle which underlies the various patterns or ornamental markings of animals--namely, that diversified coloration follows the chief lines of structure, and changes at points, such as the joints, where function changes. He says, "If we take highly decorated species--that is, animals marked by alternate dark or light bands or spots, such as the zebra, some deer, or the carnivora, we find, first, that the region of the spinal column is marked by a dark stripe; secondly, that the regions of the appendages, or limbs, are differently marked; thirdly, that the flanks are striped or spotted, along or between the regions of the lines of the ribs; fourthly, that the shoulder and hip regions are marked by curved lines; fifthly, that the pattern changes, and the direction of the lines, or spots, at the head, neck, and every joint of the limbs; and lastly, that the tips of the ears, nose, tail, and feet, and the eye are emphasised in colour. In spotted animals the greatest length of the spot is generally in the direction of the largest development of the skeleton."

This structural decoration is well seen in many insects. In caterpillars, similar spots and markings are repeated in each segment, except where modified for some form of protection. In b.u.t.terflies, the spots and bands usually have reference to the form of the wing and the arrangement of the nervures; and there is much evidence to show that the primitive markings are always spots in the cells, or between the nervures, or at the junctions of nervures, the extension and coalescence of these spots forming borders, bands, or blotches, which have become modified in infinitely varied ways for protection, warning, or recognition. Even in birds, the distribution of colours and markings follows generally the same law. The crown of the head, the throat, the ear-coverts, and the eyes have usually distinct tints in all highly coloured birds; the region of the furcula has often a distinct patch of colour, as have the pectoral muscles, the uropygium or root of the tail, and the under tail-coverts.[130]

Mr. Tylor was of opinion the primitive form of ornamentation consisted of spots, the confluence of these in certain directions forming lines or bands; and, these again, sometimes coalescing into blotches, or into more or less uniform tints covering a large portion of the surface of the body. The young lion and tiger are both spotted; and in the Java hog (Sus vittatus) very young animals are banded, but have spots over the shoulders and thighs. These spots run into stripes as the animal grows older; then the stripes expand, and at last, meeting together, the adult animal becomes of a uniform dark brown colour. So many of the species of deer are spotted when young, that Darwin concludes the ancestral form, from which all deer are derived, must have been spotted. Pigs and tapirs are banded or spotted when young; an imported young specimen of Tapirus Bairdi was covered with white spots in longitudinal rows, here and there forming short stripes.[131] Even the horse, which Darwin supposes to be descended from a striped animal, is often spotted, as in dappled horses; and great numbers show a tendency to spottiness, especially on the haunches.

Ocelli may also be developed from spots, or from bars, as pointed out by Mr. Darwin. Spots are an ordinary form of marking in disease, and these spots sometimes run together, forming blotches. There is evidence that colour markings are in some way dependent on nerve distribution. In the disease known as frontal herpes, an eruption occurs which corresponds exactly to the distribution of the ophthalmic division of the fifth cranial nerve, mapping out all its little branches even to the one which goes to the tip of the nose. In a Hindoo suffering from herpes the pigment was destroyed in the arm along the course of the ulnar nerve, with its branches along both sides of one finger and the half of another. In the leg the sciatic and scaphenous nerves were partly mapped out, giving to the patient the appearance of an anatomical diagram.[132]

These facts are very interesting, because they help to explain the general dependence of marking on structure which has been already pointed out. For, as the nerves everywhere follow the muscles, and these are attached to the various bones, we see how it happens, that the tracts in which distinct developments of colour appear, should so often be marked out by the chief divisions of the bony structure in vertebrates, and by the segments in the annulosa. There is, however, another correspondence of even greater interest and importance.

Brilliant colours usually appear just in proportion to the development of tegumentary appendages. Among birds the most brilliant colours are possessed by those which have developed frills, crests, and elongated tails like the humming-birds; immense tail-coverts like the peac.o.c.k; enormously expanded wing-feathers, as in the argus-pheasant; or magnificent plumes from the region of the coracoids in many of the birds of paradise. It is to be noted, also, that all these accessory plumes spring from parts of the body which, in other species, are distinguished by patches of colour; so that we may probably impute the development of colour and of accessory plumage to the same fundamental cause.

Among insects, the most brilliant and varied coloration occurs in the b.u.t.terflies and moths, groups in which the wing-membranes have received their greatest expansion, and whose specialisation has been carried furthest in the marvellous scaly covering which is the seat of the colour. It is suggestive, that the only other group in which functional wings are much coloured is that of the dragonflies, where the membrane is exceedingly expanded. In like manner, the colours of beetles, though greatly inferior to those of the lepidoptera, occur in a group in which the anterior pair of wings has been thickened and modified in order to protect the vital parts, and in which these wing-covers (elytra), in the course of development in the different groups, must have undergone great changes, and have been the seat of very active growth.

_The Origin of Accessory Plumes._

Mr. Darwin supposes, that these have in almost every case been developed by the preference of female birds for such males as possessed them in a higher degree than others; but this theory does not account for the fact that these plumes usually appear in a few definite parts of the body. We require some cause to initiate the development in one part rather than in another. Now, the view that colour has arisen over surfaces where muscular and nervous development is considerable, and the fact that it appears especially upon the accessory or highly developed plumes, leads us to inquire whether the same cause has not primarily determined the development of these plumes. The immense tuft of golden plumage in the best known birds of paradise (Paradisea apoda and P. minor) springs from a very small area on the side of the breast. Mr. Frank E. Beddard, who has kindly examined a specimen for me, says that "this area lies upon the pectoral muscles, and near to the point where the fibres of the muscle converge towards their attachment to the humerus. The plumes arise, therefore, close to the most powerful muscle of the body, and near to where the activities of that muscle would be at a maximum.

Furthermore, the area of attachment of the plumes is just above the point where the arteries and nerves for the supply of the pectoral muscles, and neighbouring regions, leave the interior of the body. The area of attachment of the plume is, also, as you say in your letter, just above the junction of the coracoid and sternum." Ornamental plumes of considerable size rise from the same part in many other species of paradise birds, sometimes extending laterally in front, so as to form breast shields. They also occur in many humming-birds, and in some sun-birds and honey-suckers; and in all these cases there is a wonderful amount of activity and rapid movement, indicating a surplus of vitality, which is able to manifest itself in the development of these accessory plumes.[133]

In a quite distinct set of birds, the gallinaceae, we find the ornamental plumage usually arising from very different parts, in the form of elongated tail-feathers or tail-coverts, and of ruffs or hackles from the neck. Here the wings are comparatively little used, the most constant activities depending on the legs, since the gallinaceae are pre-eminently walking, running, and scratching birds. Now the magnificent train of the peac.o.c.k--the grandest development of accessory plumes in this order--springs from an oval or circular area, about three inches in diameter, just above the base of the tail, and, therefore, situated over the lower part of the spinal column near the insertion of the powerful muscles which move the hind limbs and elevate the tail. The very frequent presence of neck-ruffs or breast-shields in the males of birds with accessory plumes may be partly due to selection, because they must serve as a protection in their mutual combats, just as does the lion's or the horse's mane. The enormously lengthened plumes of the bird of paradise and of the peac.o.c.k can, however, have no such use, but must be rather injurious than beneficial in the bird's ordinary life. The fact that they have been developed to so great an extent in a few species is an indication of such perfect adaptation to the conditions of existence, such complete success in the battle for life, that there is, in the adult male at all events, a surplus of strength, vitality, and growth-power which is able to expend itself in this way without injury.

That such is the case is shown by the great abundance of most of the species which possess these wonderful superfluities of plumage. Birds of paradise are among the commonest birds in New Guinea, and their loud voices can be often heard when the birds themselves are invisible in the depths of the forest; while Indian sportsmen have described the peafowl as being so abundant, that from twelve to fifteen hundred have been seen within an hour at one spot; and they range over the whole country from the Himalayas to Ceylon. Why, in allied species, the development of accessory plumes has taken different forms, we are unable to say, except that it may be due to that individual variability which has served as the starting-point for so much of what seems to us strange in form, or fantastic in colour, both in the animal and vegetable world.

_Development of Accessory Plumes and their Display._

If we have found a _vera causa_ for the origin of ornamental appendages of birds and other animals in a surplus of vital energy, leading to abnormal growths in those parts of the integument where muscular and nervous action are greatest, the continuous development of these appendages will result from the ordinary action of natural selection in preserving the most healthy and vigorous individuals, and the still further selective agency of s.e.xual struggle in giving to the very strongest and most energetic the parentage of the next generation. And, as all the evidence goes to show that, so far as female birds exercise any choice, it is of "the most vigorous, defiant, and mettlesome male,"

this form of s.e.xual selection will act in the same direction, and help to carry on the process of plume development to its culmination. That culmination will be reached when the excessive length or abundance of the plumes begins to be injurious to the bearer of them; and it may be this check to the further lengthening of the peac.o.c.k's train that has led to the broadening of the feathers at the ends, and the consequent production of the magnificent eye-spots which now form its crowning ornament.

The display of these plumes will result from the same causes which led to their production. Just in proportion as the feathers themselves increased in length and abundance, the skin-muscles which serve to elevate them would increase also; and the nervous development as well as the supply of blood to these parts being at a maximum, the erection of the plumes would become a habit at all periods of nervous or s.e.xual excitement. The display of the plumes, like the existence of the plumes themselves, would be the chief external indication of the maturity and vigour of the male, and would, therefore, be necessarily attractive to the female. We have, thus, no reason for imputing to her any of those aesthetic emotions which are excited in us, by the beauty of form, colour, and pattern of these plumes; or the still more improbable aesthetic tastes, which would cause her to choose her mate on account of minute differences in their forms, colours, or patterns.

As co-operating causes in the production of accessory ornamental plumes, I have elsewhere suggested[134] that crests and other erectile feathers may have been useful in making the bird more formidable in appearance, and thus serving to frighten away enemies; while long tail or wing feathers might serve to distract the aim of a bird of prey. But though this might be of some use in the earlier stages of their development, it is probably of little importance compared with the vigour and pugnacity of which the plumes are the indication, and which enable most of their possessors to defend themselves against the enemies which are dangerous to weaker and more timid birds. Even the tiny humming-birds are said to attack birds of prey that approach too near to their nests.

_The Effect of Female Preference will be Neutralised by Natural Selection._

The various facts and arguments now briefly set forth, afford an explanation of the phenomena of male ornament, as being due to the general laws of growth and development, and make it unnecessary to call to our aid so hypothetical a cause as the c.u.mulative action of female preference. There remains, however, a general argument, arising from the action of natural selection itself, which renders it almost inconceivable that female preference could have been effective in the way suggested; while the same argument strongly supports the view here set forth. Natural selection, as we have seen in our earlier chapters, acts perpetually and on an enormous scale in weeding out the "unfit" at every stage of existence, and preserving only those which are in all respects the very best. Each year, only a small percentage of young birds survive to take the place of the old birds which die; and the survivors will be those which are best able to maintain existence from the egg onwards, an important factor being that their parents should be well able to feed and protect them, while they themselves must in turn be equally able to feed and protect their own offspring. Now this extremely rigid action of natural selection must render any attempt to select mere ornament utterly nugatory, unless the most ornamented always coincide with "the fittest" in every other respect; while, if they do so coincide, then any selection of ornament is altogether superfluous. If the most brightly coloured and fullest plumaged males are _not_ the most healthy and vigorous, have _not_ the best instincts for the proper construction and concealment of the nest, and for the care and protection of the young, they are certainly not the fittest, and will not survive, or be the parents of survivors. If, on the other hand, there _is_ generally this correlation--if, as has been here argued, ornament is the natural product and direct outcome of superabundant health and vigour, then no other mode of selection is needed to account for the presence of such ornament. The action of natural selection does not indeed disprove the existence of female selection of ornament as ornament, but it renders it entirely ineffective; and as the direct evidence for any such female selection is almost _nil_, while the objections to it are certainly weighty, there can be no longer any reason for upholding a theory which was provisionally useful in calling attention to a most curious and suggestive body of facts, but which is now no longer tenable. The term "s.e.xual selection" must, therefore, be restricted to the direct results of male struggle and combat. This is really a form of natural selection, and is a matter of direct observation; while its results are as clearly deducible as those of any of the other modes in which selection acts. And if this restriction of the term is needful in the case of the higher animals it is much more so with the lower. In b.u.t.terflies the weeding out by natural selection takes place to an enormous extent in the egg, larva, and pupa states; and perhaps not more than one in a hundred of the eggs laid produces a perfect insect which lives to breed. Here, then, the impotence of female selection, if it exist, must be complete; for, unless the most brilliantly coloured males are those which produce the best protected eggs, larvae, and pupae, and unless the particular eggs, larvae, and pupae, which are able to survive, are those which produce the most brilliantly coloured b.u.t.terflies, any choice the female might make must be completely swamped. If, on the other hand, there _is_ this correlation between colour development and perfect adaptation to conditions in all stages, then this development will necessarily proceed by the agency of natural selection and the general laws which determine the production of colour and of ornamental appendages.[135]

_General Laws of Animal Coloration._

The condensed account which has now been given of the phenomena of colour in the animal world will sufficiently show the wonderful complexity and extreme interest of the subject; while it affords an admirable ill.u.s.tration of the importance of the great principle of utility, and of the effect of the theories of natural selection and development in giving a new interest to the most familiar facts of nature. Much yet remains to be done, both in the observation of new facts as to the relations between the colours of animals and their habits or economy, and, more especially, in the elucidation of the laws of growth which determine changes of colour in the various groups; but so much is already known that we are able, with some confidence, to formulate the general principles which have brought about all the beauty and variety of colour which everywhere delight us in our contemplation of animated nature. A brief statement of these principles will fitly conclude our exposition of the subject.

1. Colour may be looked upon as a necessary result of the highly complex chemical const.i.tution of animal tissues and fluids. The blood, the bile, the bones, the fat, and other tissues have characteristic, and often brilliant colours, which we cannot suppose to have been determined for any special purpose, as colours, since they are usually concealed. The external organs, with their various appendages and integuments, would, by the same general laws, naturally give rise to a greater variety of colour.

2. We find it to be the fact that colour increases in variety and intensity as external structures and dermal appendages become more differentiated and developed. It is on scales, hair, and especially on the more highly specialised feathers, that colour is most varied and beautiful; while among insects colour is most fully developed in those whose wing membranes are most expanded, and, as in the lepidoptera, are clothed with highly specialised scales. Here, too, we find an additional mode of colour production in transparent lamellae or in fine surface striae which, by the laws of interference, produce the wonderful metallic hues of so many birds and insects.

3. There are indications of a progressive change of colour, perhaps in some definite order, accompanying the development of tissues or appendages. Thus spots spread and fuse into bands, and when a lateral or centrifugal expansion has occurred--as in the termination of the peac.o.c.ks' train feathers, the outer web of the secondary quills of the Argus pheasant, or the broad and rounded wings of many b.u.t.terflies--into variously shaded or coloured ocelli. The fact that we find gradations of colour in many of the more extensive groups, from comparatively dull or simple to brilliant and varied hues, is an indication of some such law of development, due probably to progressive local segregation in the tissues of identical chemical or organic molecules, and dependent on laws of growth yet to be investigated.

4. The colours thus produced, and subject to much individual variation, have been modified in innumerable ways for the benefit of each species.

The most general modification has been in such directions as to favour concealment when at rest in the usual surroundings of the species, sometimes carried on by successive steps till it has resulted in the most minute imitation of some inanimate object or exact mimicry of some other animal. In other cases bright colours or striking contrasts have been preserved, to serve as a warning of inedibility or of dangerous powers of attack. Most frequent of all has been the specialisation of each distinct form by some tint or marking for purposes of easy recognition, especially in the case of gregarious animals whose safety largely depends upon a.s.sociation and mutual defence.

5. As a general rule the colours of the two s.e.xes are alike; but in the higher animals there appears a tendency to deeper or more intense colouring in the male, due probably to his greater vigour and excitability. In many groups in which this superabundant vitality is at a maximum, the development of dermal appendages and brilliant colours has gone on increasing till it has resulted in a great diversity between the s.e.xes; and in most of these cases there is evidence to show that natural selection has caused the female to retain the primitive and more sober colours of the group for purposes of protection.

_Concluding Remarks._

The general principles of colour development now sketched out enable us to give some rational explanation of the wonderful amount of brilliant colour which occurs among tropical animals. Looking on colour as a normal product of organisation, which has either been allowed free play, or has been checked and modified for the benefit of the species, we can see at once that the luxuriant and perennial vegetation of the tropics, by affording much more constant means of concealment, has rendered brilliant colour less hurtful there than in the temperate and colder regions. Again, this perennial vegetation supplies abundance of both vegetable and insect food throughout the year, and thus a greater abundance and greater variety of the forms of life are rendered possible, than where recurrent seasons of cold and scarcity reduce the possibilities of life to a minimum. Geology furnishes us with another reason, in the fact, that throughout the tertiary period tropical conditions prevailed far into the temperate regions, so that the possibilities of colour development were still greater than they are at the present time. The tropics, therefore, present to us the results of animal development in a much larger area and under more favourable conditions than prevail to-day. We see in them samples of the productions of an earlier and a better world, from an animal point of view; and this probably gives a greater variety and a finer display of colour than would have been produced, had conditions always been what they are now. The temperate zones, on the other hand, have recently suffered the effects of a glacial period of extreme severity, with the result that almost the only gay coloured birds they now possess are summer visitors from tropical or sub-tropical lands. It is to the unbroken and almost unchecked course of development from remote geological times that has prevailed in the tropics, favoured by abundant food and perennial shelter, that we owe such superb developments as the frills and crests and jewelled shields of the humming-birds, the golden plumes of the birds of paradise, and the resplendent train of the peac.o.c.k. This last exhibits to us the culmination of that marvel and mystery of animal colour which is so well expressed by a poet-artist in the following lines. The marvel will ever remain to the sympathetic student of nature, but I venture to hope that in the preceding chapters I have succeeded in lifting--if only by one of its corners--the veil of mystery which has for long shrouded this department of nature.

_On a Peac.o.c.k's Feather._

In Nature's workshop but a shaving, Of her poem but a word, But a tint brushed from her palette, This feather of a bird!

Yet set it in the sun glance, Display it in the shine, Take graver's lens, explore it, Note filament and line, Mark amethyst to sapphire, And sapphire to gold, And gold to emerald changing The archetype unfold!

Tone, tint, thread, tissue, texture, Through every atom scan, Conforming still, developing, Obedient to plan.

This but to form a pattern On the garment of a bird!

What then must be the poem, This but its lightest word!

Sit before it; ponder o'er it, 'Twill thy mind advantage more, Than a treatise, than a sermon, Than a library of lore.

FOOTNOTES:

[Footnote 118: Darwin's _Descent of Man_, p. 271.]

[Footnote 119: Darwin's _Descent of Man_, p. 294, and footnote.]

[Footnote 120: _Nature_, 1871, p. 489.]

[Footnote 121: Darwin in _Nature_, 1880, p. 237.]

[Footnote 122: See the author's _Contributions to Natural Selection_, chap. vii. in which these facts were first brought forward.]

[Footnote 123: On this point see the author's _Contributions to Natural Selection_, chap. v. i.]

[Footnote 124: Seebohm's _History of British Birds_, vol. ii., introduction, p. xiii.]

[Footnote 125: For details see Darwin's _Descent of Man_, chap. xii.]

[Footnote 126: _Descent of Man_, pp. 417, 418, 420.]