Part 9

See Macmillan’s Magazine, Sept. 1867, “On the Colours of Animals and Plants,” and Chapters V. and VI. of the present volume.

The amount of variation which is likely to occur in a species will be greatly influenced by two factors--the occurrence of a change in the physical conditions, and the average abundance or scarcity of the individuals composing the species. When from these or other causes variation occurs, it may become fixed as a variety or a race, or may go on increasing to a certain extent, either from a tendency to vary along certain special lines induced by local or physiological causes, or by the continued survival and propagation of all such varieties as are beneficial to the race. After a certain time a balance will be arrived at, either by the limits of useful variation in this one direction having been reached, or by the species becoming harmoniously adapted to all the surrounding conditions; and without some change in these conditions the specific form may then remain unaltered for a very long time; whence arises the common impression of the fixity of species. Now in a country like Chili, forming part of a great continent very well stocked with all forms of organic life, the majority of the species would be in a state of stable equilibrium; the most favourable variations would have been long ago selected; and the numbers of individuals in each species would be tolerably constant, being limited by the numerous other forms whose food and habits were similar, or which in any way impinged upon its sphere of existence. We may, therefore, assume that the Chilian humming-bird which migrated to Juan Fernandez was a stable form, hardly if at all different from the existing species which is termed Eustephanus galeritus. On the island it met with very changed but highly favourable conditions,--an abundant shrubby vegetation and a tolerably rich flora; less extremes of climate than on the mainland; and, most important of all, absolute freedom from the competition of rival species. The flowers and their insect inhabitants were all its own; there were no snakes or mammalia to plunder its nests; nothing to prevent the full enjoyment of existence. The consequence would be, rapid increase and a large permanent population, which still maintains itself; for Mr. Moseley, of the Challenger expedition, has informed the writer that humming-birds are extraordinarily abundant in Juan Fernandez, every bush or tree having one or two darting about it. Here, then, we have one of the special conditions which have always been held to favour variation--a great increase in the number of individuals; but, as there was no struggle with allied creatures, there was no need for any modification in form or structure, and we accordingly find that the only important variations which have become permanent are those of size and of colour. The increased size would naturally arise from greater abundance of food with a more equable climate throughout the year, the healthier, stronger, and larger individuals being preserved. The change of colour would depend on molecular changes in the plumage accompanying the increase of size; and the superior energy and vitality in the male, aided by the favourable change in conditions and rapid increase of population, would lead to an increased intensity of colour, the special tint being determined either by local conditions or by inherited tendencies in the race. It is to be noted that the change from green to red is in the direction of the less refrangible rays of the spectrum, and is in accordance with the law of change which has been shown to accompany expansion in inorganic,--growth and development in organic forms. The change of colour in the female, not being urged on by such intense vital activity as in the case of the male, would be much slower, and, owing probably to inherited tendencies, in a different direction. The under-surface of the Chilian bird is ashy with bronzy-green spots on the breast, while the tail is entirely bronze-green. In the Juan Fernandez species the under-surface has become pure white, the breast-spots larger and of a purer golden-green, while the whole inner web of the tail-feathers has become pure white, producing a most elegant effect when the tail is expanded.

See “Colours of Animals,” Macmillan’s Magazine, Sept. 1877, pp. 394-398, and Chapter V. in the present volume.

We may now follow the two sexes to the remoter island, at a period when the male had acquired his permanent style of colouring, but was not quite so large as he subsequently became; while the change of the female bird had not been half completed. In this small and comparatively barren island (a mere rock, as it is described by some authors) there would be no such constant abundance of food, and therefore no possibility of a large permanent population; while the climate would not differ materially from that of the larger island. Variation would therefore be checked, or might be stopped altogether; and we find the facts exactly correspond to this view. The male, which had already acquired his colour, remains almost undistinguishable from his immediate ancestral form; but he is a little smaller, indicating either that the full size of that form had not been acquired at the period of migration, or that a slight diminution of size has since occurred, owing to a deficiency of food. The female shows also a slight diminution of size, but in other respects is almost exactly intermediate between the Chilian and Juan Fernandez females. The colour beneath is light ashy, the breast-spots are intermediate in size and colour, and the tail-feathers have a large ill-defined white spot on the end of the inner web which has only to be extended along the whole web to produce the exact character which has been acquired in Juan Fernandez. It seems probable, therefore, that the female bird has remained nearly or quite stationary since its migration, while its Juan Fernandez relative has gone on steadily changing in the direction already begun; and the more distant species geographically thus appears to be more nearly related to its Chilian ancestor.

Coming down to a more recent period, we find that the comparatively small and dull-coloured Chilian bird has again migrated to Juan Fernandez; but it at once came into competition with its red descendant, which had firm possession of the soil, and had probably undergone slight constitutional changes exactly fitting it to its insular abode. The new-comer, accordingly, only just manages to maintain its footing; for we are told by Mr. Reed, of Santiago, that it is by no means common; whereas, as we have seen, the red species is excessively abundant. We may further suspect that the Chilian birds now pass over pretty frequently to Juan Fernandez, and thus keep up the stock; for it must be remembered that whereas, at a first migration, both a male and a female are necessary for colonization, yet, after a colony is formed, any stray bird which may come over adds to the numbers, and checks permanent variation by cross-breeding.

We find, then, that all the chief peculiarities of the three allied species of humming-birds which inhabit the Juan Fernandez group of islands, may be fairly traced to the action of those general laws which Mr. Darwin and others have shown to determine the variations of animals and the perpetuation of those variations. It is also instructive to note, that where the variations of colour and size have been greatest they are accompanied by several lesser variations in other characters. In the Juan Fernandez bird the bill has become a little shorter, the tail-feathers somewhat broader, and the fiery cap on the head somewhat smaller; all these peculiarities being less developed or absent in the birds inhabiting Mas-afuera. These coincident changes may be due, either to what Mr. Darwin has termed correlation of growth, or to the partial reappearance of ancestral characters under more favourable conditions, or to the direct action of changes of climate and of food; but they show us how varied and unaccountable are the changes in specific forms that may be effected in a comparatively short time, and by means of very slight changes of locality.

If now we consider the enormously varied conditions presented by the whole continent of America--the hot, moist, and uniform forest-plains of the Amazon; the open llanos of the Orinoco; the dry uplands of Brazil; the sheltered valleys and forest slopes of the Eastern Andes; the verdant plateaus, the barren paramos, the countless volcanic cones with their peculiar Alpine vegetation; the contrasts of the East and West coasts; the isolation of the West Indian islands, and to a less extent of Central America and Mexico which we know have been several times separated from South America; and when we further consider that all these characteristically distinct areas have been subject to cosmical and local changes, to elevations and depressions, to diminution and increase of size, to greater extremes and greater uniformity of temperature, to increase or decrease of rainfall; and that with these changes there have been coincident changes of vegetation and of animal life, all affecting in countless ways the growth and development, the forms and colours, of these wonderful little birds--if we consider all these varied and complex influences, we shall be less surprised at their strange forms, their infinite variety, their wondrous beauty. For how many ages the causes above enumerated may have acted upon them we cannot say; but their extreme isolation from all other birds, no less than the abundance and variety of their generic and specific forms, clearly point to a very high antiquity.

The Relations and Affinities of Humming-birds.--The question of the position of this family in the class of birds and its affinities or resemblances to other groups, is so interesting, and affords such good opportunities for explaining some of the best-established principles of classification in natural history in a popular way, that we propose to discuss it at some length, but without entering into technical details.

There is in the Eastern hemisphere, especially in tropical Africa and Asia, a family of small birds called Sun-birds, which are adorned with brilliant metallic colours, and which, in shape and general appearance, much resemble humming-birds. They frequent flowers in the same way, feeding on honey and insects; and all the older naturalists placed the two families side by side as undoubtedly allied. In the year 1850, in a general catalogue of birds, Prince Lucien Bonaparte, a learned ornithologist, placed the humming-birds next to the swifts, and far removed from the Nectarinidæ or sun-birds; and this view of their position has gained ground with increasing knowledge, so that now all the more advanced ornithologists have adopted it. Before proceeding to point out the reasons for this change of view, it will be well to discuss a few of the general principles which guide naturalists in the solution of such problems.

How to Determine Doubtful Affinities.--It is now generally admitted that, for the purpose of determining obscure and doubtful affinities, we must examine by preference those parts of an animal which have little or no direct influence on its habits and general economy. The value of an organ, or of any detail of structure, for purposes of classification, is generally in inverse proportion to its adaptability to special uses. And the reason of this is apparent, when we consider that similarities of food and habits are often accompanied by similarities of external form or of special organs, in totally distinct animals. Porpoises, for example, are modified externally so as to resemble fishes; yet they are really mammalia. Some marsupials are carnivorous, and are so like true carnivora that it is only by minute peculiarities of structure that the skeleton of the one can be distinguished from that of the other. Many of the hornbills and toucans have the same general form, and resemble each other in habits, in food, and in their enormous bills; yet peculiarities in the structure of the feet, in the form of the breast-bone, in the cranium, and in the texture and arrangement of the plumage, show that they have no real affinity, the former approaching the kingfishers, the latter the cuckoos. Such structural peculiarities as these have no direct relation to habits; and they are therefore little liable to change, when from any cause a portion of the group may have been driven to adopt a new mode of life. Thus all the Old World apes, however much they may differ in size or habits, and whether we class them as baboons, monkeys, or gorillas, have the same number of teeth; while the American monkeys all have an additional premolar tooth. This difference can have no relation to the habits of the two groups, because each group exhibits differences of habits greater than often occur between American and Asiatic species; and it thus becomes a valuable character indicating the radical distinctness of the two groups, a distinctness confirmed by other anatomical characters.

On the other hand, peculiarities of organization which seem specially adapted to certain modes of life, are often diminished or altogether lost in a few species of the group, showing their essential unimportance to the type, as well as their small value for classification. Thus, the woodpeckers are most strikingly characterised by a very long and highly extensible tongue, with the muscles attached to the tongue-bone prolonged backward over the head so as to enable the tongue to be suddenly darted out; and also by the rigid and pointed tail which is a great help in climbing up the vertical trunks of trees. But in one group (the Picumni), the tail becomes quite soft, while the tongue remains fully developed; and in another (Meiglyptes) the characteristic tail remains, while the prolonged hyoid muscles have almost entirely disappeared, and the tongue has consequently lost its peculiar extensile power; yet in both these cases the form of the breast-bone and the character of the feet, the skeleton, and the plumage, show that the birds are really woodpeckers; while even the habits and the food are very little altered. In like manner the bill may undergo great changes; as from the short crow-like bill of the true birds-of-paradise to the long slender bills of Epimachinæ, which latter were on that account long classed apart in the tribe of Tenuirostres, or slender-billed birds, but whose entire structure shows them to be closely allied to the paradise-birds. So, the long feathery tongue of the toucans differs from that of every other bird; yet it is not held to overbalance the weight of anatomical peculiarities which show that these birds are allied to the barbets and the cuckoos.

The skeleton, therefore, and especially the sternum or breast-bone, affords us an almost infallible guide in doubtful cases; because it appears to change its form with extreme slowness, and thus indicates deeper-seated affinities than those shown by organs which are in direct connection with the outside world, and are readily modified in accordance with varying conditions of existence. Another, though less valuable guide is afforded, in the case of birds, by the eggs. These often have a characteristic form and colour, and a peculiar texture of surface, running unchanged through whole genera and families which are nearly related to each other, however much they may differ in outward form and habits. Another detail of structure which has no direct connection with habits and economy, is the manner in which the plumage is arranged on the body. The feathers of birds are by no means set uniformly over their skin, but grow in certain definite lines and patches, which vary considerably in shape and size in the more important orders and tribes, while the mode of arrangement agrees in all which are known to be closely related to each other; and thus the form of the feather-tracts or the “pterylography” as it is termed, of a bird, is a valuable aid in doubtful cases of affinity.

Now, if we apply these three tests to the humming-birds, we find them all pointing in the same direction. The sternum or breast-bone is not notched behind; and this agrees with the swifts, and not with the sun-birds, whose sternum has two deep notches behind, as in all the families of the vast order of Passeres to which the latter belong. The eggs of both swifts and humming-birds are white, only two in number, and resembling each other in texture. And in the arrangement of the feather-tracts the humming-birds approach more nearly to the swifts than they do to any other birds; and altogether differ from the sun-birds, which, in this respect as in so many others, resemble the honey-suckers of Australia and other true passerine birds.

Resemblances of Swifts and Humming-birds.--Having this clue to their affinities, we shall find other peculiarities common to these two groups, the swifts and the humming-birds. They have both ten tail-feathers, while the sun-birds have twelve. They have both only sixteen true quill-feathers, and they are the only birds which have so small a number. The humming-birds are remarkable for having, in almost all the species, the first quill the longest of all, the only other birds resembling them in this respect being a few species of swifts; and, lastly, in both groups the plumage is remarkably compact and closely pressed to the body. Yet, with all these points of agreement, we find an extreme diversity in the bills and tongues of the two groups. The swifts have a short, broad, flat bill, with a flat horny-tipped tongue of the usual character; while the humming-birds have a very long, narrow, almost cylindrical bill, containing a tubular and highly extensible tongue. The essential point however is, that whereas hardly any of the other characters we have adduced are adaptive, or strictly correlated with habits and economy, this character is pre-eminently so; for the swifts are pure aërial insect-hunters, and their short, broad bills, and wide gape, are essential to their mode of life. The humming-birds, on the other hand, are floral insect-hunters, and for this purpose their peculiarly long bills and extensile tongues are especially adapted; while they are at the same time honey-suckers, and for this purpose have acquired the tubular tongue. The formation of such a tubular tongue out of one of the ordinary kind is easily conceivable, as it only requires to be lengthened, and the two laminæ of which it is composed curled in at the sides; and these changes it probably goes through in the young birds.

When on the Amazon I once had a nest brought me containing two little unfledged humming-birds, apparently not long hatched. Their beaks were not at all like those of their parents, but short, triangular, and broad at the base; just the form of the beak of a swallow or swift slightly lengthened. Thinking (erroneously) that the young birds were fed by their parents on honey, I tried to feed them with a syrup made of honey and water, but though they kept their mouths constantly open as if ravenously hungry, they would not swallow the liquid, but threw it out again and sometimes nearly choked themselves in the effort. At length I caught some minute flies, and on dropping one of these into the open mouth it instantly closed, the fly was gulped down and the mouth opened again for more; and each took in this way fifteen or twenty little flies in succession before it was satisfied. They lived thus three or four days, but required more constant care than I could give them. These little birds were in the “swift” stage; they were pure insect-eaters, with a bill and mouth adapted for insect-eating only. At that time I was not aware of the importance of the observation of the tongue; but as the bill was so short and the tubular tongue not required, there can be little doubt that the organ was, at that early stage of growth, short and flat, as it is in the birds most nearly allied to them.

Differences between Sun-birds and Humming-birds.--In respect of all the essential and deep-seated points of structure, which have been shown to offer such remarkable similarities between the swifts and the humming-birds, the sun-birds of the Eastern hemisphere differ totally from the latter, while they agree with the passerine birds generally, or more particularly with the creepers and honey-suckers. They have a deeply-notched sternum; they have twelve tail-feathers in place of ten; they have nineteen quills in place of sixteen; and the first quill instead of being the longest is the very shortest of all, while the wings are short and round, instead of being excessively long and pointed; their plumage is arranged differently; and their feet are long and strong, instead of being excessively short and weak. There remain only the superficial characters of small size and brilliant metallic colours to assimilate them with the humming-birds, and one structural feature--a tubular and somewhat extensile tongue. This, however, is a strictly adaptive character, the sun-birds feeding on small insects and the nectar of flowers, just as do the humming-birds; and it is a remarkable instance of a highly peculiar modification of an organ occurring independently in two widely-separate groups. In the sun-birds the hyoid or tongue-muscles do not extend so completely over the head as they do in the humming-birds, so that the tongue is less extensible; but it is constructed in exactly the same way by the inrolling of the two laminæ of which it is composed.

The tubular tongue of the sun-birds is a special adaptive modification acquired within the family itself, and not inherited from a remote ancestral form. This is shown by the amount of variation this organ exhibits in different members of the family. It is most highly developed in the Arachnotheræ, or spider-hunters, of Asia, which are sun-birds without any metallic or other brilliant colouring. These have the longest bills and tongues, and the most developed hyoid muscles; they hunt much about the blossoms of palm-trees, and may frequently be seen probing the flowers while fluttering clumsily in the air, just as if they had seen and attempted to imitate the aërial gambols of the American humming-birds. The true metallic sun-birds generally cling about the flowers with their strong feet; and they feed chiefly on minute hard insects, as do many humming-birds. There is, however, one species (Chalcoparia phœnicotis) always classed as a sun-bird, which differs entirely from the rest of the species in having the tongue flat, horny, and forked at the tip; and its food seems to differ correspondingly, for small caterpillars were found in its stomach. More remotely allied, but yet belonging to the same family, are the little flower-peckers of the genus Diceum, which have a short bill and a tongue twice split at the end; and these feed on small fruits, and perhaps on buds and on the pollen of flowers. The little white-eyes (Zosterops), which are probably allied to the last, eat soft fruits and minute insects.

Here then we have an extensive group of birds, considerably varied in external form, yet undoubtedly closely allied to each other, one division of which is specially adapted to feed on the juices secreted by flowers and the minute insects that harbour in them; and these alone have a lengthened bill and double tubular tongue, just as in the humming-birds. We can hardly have a more striking example of the necessity of discriminating between adaptive and purely structural characters. The same adaptive character may coexist in two groups which have a similar mode of life, without indicating any affinity between them, because it may have been acquired by each independently, to enable it to fill a similar place in nature. In such cases it is found to be an almost isolated character, apparently connecting two groups which otherwise differ radically. Non-adaptive, or purely structural characters, on the other hand, are such as have probably been transmitted from a remote ancestor; and thus indicate fundamental peculiarities of growth and development. The changes of structure rendered necessary by modifications of the habits or instincts of the different species, have been made, to a great extent, independently of such characters; and as several of these may always be found in the same animal their value becomes cumulative. We thus arrive at the seeming paradox, that the less of direct use is apparent in any peculiarity of structure, the greater is its value in indicating true, though perhaps remote, affinities; while any peculiarity of an organ which seems essential to its possessor’s well-being is often of very little value in indicating its affinity for other creatures.

This somewhat technical discussion will, it is hoped, enable the general reader to understand some of the more important principles of the modern or natural classification of animals, as distinguished from the artificial system which long prevailed. It will also afford him an easily remembered example of those principles, in the radical distinctness of two families of birds often confounded together,--the sun-birds of the Eastern Hemisphere, and the humming-birds of America; and in the interesting fact that the latter are essentially swifts--profoundly modified, it is true, for an aërial and flower-haunting existence, but still bearing in many important peculiarities of structure the unmistakable evidences of a common origin.

THE COLOURS OF ANIMALS AND SEXUAL SELECTION.

General Phenomena of Colour in the Organic World--Theory of Heat and Light as producing Colour--Changes of Colour in Animals produced by Coloured Light--Classification of Organic Colours--Protective Colour--Warning Colours--Sexual Colours--Typical Colours--The Nature of Colour--Colour a normal product of Organization--Theory of Protective Colours--Theory of Warning Colours--Theory of Sexual Colours--Colour as a means of Recognition--Colour proportionate to Integumentary Development--Selection by Females not a cause of Colour--Probable use of the Horns of Beetles--Cause of the greater brilliancy of some Female Insects--Theory of display of Ornaments by Males--Natural Selection as neutralizing Sexual Selection--Theory of Typical Colours--Colour-development as illustrated by Humming-birds--Local causes of Colour-development--Summary on Colour-development in Animals.

There is probably no one quality of natural objects from which we derive so much pure and intellectual enjoyment as from their colours. The heavenly blue of the firmament, the glowing tints of sunset, the exquisite purity of the snowy mountains, and the endless shades of green presented by the verdure-clad surface of the earth, are a never-failing source of pleasure to all who enjoy the inestimable gift of sight. Yet these constitute, as it were, but the frame and background of a marvellous and ever-changing picture. In contrast with these broad and soothing tints, we have presented to us in the vegetable and animal worlds, an infinite variety of objects adorned with the most beautiful and most varied hues. Flowers, insects and birds, are the organisms most generally ornamented in this way; and their symmetry of form, their variety of structure, and the lavish abundance with which they clothe and enliven the earth, cause them to be objects of universal admiration. The relation of this wealth of colour to our mental and moral nature is indisputable. The child and the savage alike admire the gay tints of flower, bird, and insect; while to many of us their contemplation brings a solace and enjoyment which is both intellectually, and morally beneficial. It can then hardly excite surprise that this relation was long thought to afford a sufficient explanation of the phenomena of colour in nature; and although the fact that--

“Full many a flower is born to blush unseen, And waste its sweetness on the desert air--”

might seem to throw some doubt on the sufficiency of the explanation, the answer was easy,--that in the progress of discovery, man would, sooner or later, find out and enjoy every beauty that the hidden recesses of the earth have in store for him. This theory received great support, from the difficulty of conceiving any other use or meaning in the colours with which so many natural objects are adorned. Why should the homely gorse be clothed in golden raiment, and the prickly cactus be adorned with crimson bells? Why should our fields be gay with buttercups, and the heather-clad mountains be clad in purple robes? Why should every land produce its own peculiar floral gems, and the alpine rocks glow with beauty, if not for the contemplation and enjoyment of man? What could be the use to the butterfly of its gaily-painted wings, or to the humming-bird of its jewelled breast, except to add the final touches to a world-picture, calculated at once to please and to refine mankind? And even now, with all our recently-acquired knowledge of this subject, who shall say that these old-world views were not intrinsically and fundamentally sound; and that, although we now know that colour has “uses” in nature that we little dreamt of, yet the relation of those colours--or rather of the various rays of light--to our senses and emotions, may not be another, and perhaps more important use which they subserve in the great system of the universe?

We now propose to lay before our readers a general account of the more recent discoveries on this interesting subject; and in doing so, it will be necessary first to give an outline of the more important facts as to the colours of organised beings; then to point out the cases in which it has been shown that colour is of use; and lastly, to endeavour to throw some light on its nature, and on the general laws of its development.

Among naturalists, colour was long thought to be of little import, and to be quite untrustworthy as a specific character. The numerous cases of variability of colour led to this view. The occurrence of white blackbirds, white peacocks, and black leopards; of white blue-bells, and of white, blue, or pink milkworts, led to the belief that colour was essentially unstable, that it could therefore be of little or no importance, and belonged to quite a different class of characters from form or structure. But it now begins to be perceived that these cases, though tolerably numerous, are, after all, exceptional; and that colour, as a rule, is a constant character. The great majority of species, both of animals and plants, are each distinguished by peculiar tints which vary very little, while the minutest markings are often constant in thousands or millions of individuals. All our field buttercups are invariably yellow, and our poppies red; while many of our butterflies and birds resemble each other in every spot and streak of colour through thousands of individuals. We also find that colour is constant in whole genera and other groups of species. The Genistas are all yellow, the Erythrinas all red; many genera of Carabidæ are entirely black; whole families of birds--as the Dendrocolaptidæ--are brown; while among butterflies the numerous species of Lycæna are all more or less blue, those of Pontia white, and those of Callidryas yellow. An extensive survey of the organic world thus leads us to the conclusion that colour is by no means so unimportant or inconstant a character as at first sight it appears to be; and the more we examine it the more convinced we shall become that it must serve some purpose in nature, and that, besides charming us by its diversity and beauty, it must be well worthy of our attentive study, and have many secrets to unfold to us.

Theory of Heat and Light as producing Colour.--In commencing our study of the great mass of facts relating to the colours of the organic world, it will be necessary to consider first, how far the chief theories already proposed will account for them. One of the most obvious and most popular of these theories, and one which is still held, in part at least, by many eminent naturalists, is--that colour is due to some direct action of the heat and light of the sun--thus at once accounting for the great number of brilliant birds, insects, and flowers, which are found between the tropics.

But before proceeding to discuss this supposed explanation of the colours of living things we must ask the preliminary question,--whether it is really the fact that colour is more developed in tropical than in temperate climates, in proportion to the whole number of species; and even if we find this to be so, we have to inquire whether there are not so many and such striking exceptions to the rule, as to indicate some other causes at work than the direct influence of solar light and heat. As this is a most important branch of the inquiry, we must go into it somewhat fully.