Let us then suppose that a given species consists of 100,000 individuals of each sex, with only the usual amount of fluctuating external variability. Let a physiological variation arise, so that 10 per cent of the whole number—10,000 individuals of each sex—while remaining fertile inter se become quite sterile with the remaining 90,000. This peculiarity is not correlated with any external differences of form or colour, or with inherent peculiarities of likes or dislikes leading to any choice as to the pairing of the two sets of individuals. We have now to inquire, What would be the result?
Taking, first, the 10,000 pairs of the physiological or abnormal variety, we find that each male of these might pair with any one of the whole 100,000 of the opposite sex. If, therefore, there was nothing to limit their choice to particular individuals of either variety, the probabilities are that 9000 of them would pair with the opposite variety, and only 1000 with their own variety—that is, that 9000 would form sterile unions, and only one thousand would form fertile unions.
Taking, next, the 90,000 normal individuals of either sex, we find, that each male of these has also a choice of 100,000 to pair with. The probabilities are, therefore, that nine-tenths of them—that is, 81,000—would pair with their normal fellows, while 9000 would pair with the opposite abnormal variety forming the above-mentioned sterile unions.
Now, as the number of individuals forming a species remains constant, generally speaking, from year to year, we shall have next year also 100,000 pairs, of which the two physiological varieties will be in the proportion of eighty-one to one, or 98,780 pairs of the normal variety to 1220[64] of the abnormal, that being the proportion of the fertile unions of each. In this year we shall find, by the same rule of probabilities, that only 15 males of the abnormal variety will pair with their like and be fertile, the remaining 1205 forming sterile unions with some of the normal variety. The following year the total 100,000 pairs will consist of 99,984 of the normal, and only 16 of the abnormal variety; and the probabilities, of course, are, that the whole of these latter will pair with some of the enormous preponderance of normal individuals, and, their unions being sterile, the physiological variety will become extinct in the third year.
If now in the second and each succeeding year a similar proportion as at first (10 per cent) of the physiological variety is produced afresh from the ranks of the normal variety, the same rate of diminution will go on, and it will be found that, on the most favourable estimate, the physiological variety can never exceed 12,000 to the 88,000 of the normal form of the species, as shown by the following table:—
2d " 1,220 + 10,000 again produced.
3d " 16 + 1,220 + 10,000 do. = 11,236
4th " O + 16 + 1,220 + 10,000 do. = 11,236
5th " O + 16 + 1,220 + 10,000 = 11,236
and so on for any number of generations.
In the preceding discussion we have given the theory the advantage of the large proportion of 10 per cent of this very exceptional variety arising in its midst year by year, and we have seen that, even under these favourable conditions, it is unable to increase its numbers much above its starting-point, and that it remains wholly dependent on the continued renewal of the variety for its existence beyond a few years. It appears, then, that this form of inter-specific sterility cannot be increased by natural or any other known form of selection, but that it contains within itself its own principle of destruction. If it is proposed to get over the difficulty by postulating a larger percentage of the variety annually arising within the species, we shall not affect the law of decrease until we approach equality in the numbers of the two varieties. But with any such increase of the physiological variety the species itself would inevitably suffer by the large proportion of sterile unions in its midst, and would thus be at a great disadvantage in competition with other species which were fertile throughout. Thus, natural selection will always tend to weed out any species with too great a tendency to sterility among its own members, and will therefore prevent such sterility from becoming the general characteristic of varying species, which this theory demands should be the case.
On the whole, then, it appears clear that no form of infertility or sterility between the individuals of a species, can be increased by natural selection unless correlated with some useful variation, while all infertility not so correlated has a constant tendency to effect its own elimination. But the opposite property, fertility, is of vital importance to every species, and gives the offspring of the individuals which possess it, in consequence of their superior numbers, a greater chance of survival in the battle of life. It is, therefore, directly under the control of natural selection, which acts both by the self-preservation of fertile and the self-destruction of infertile stocks—except always where correlated as above, when they become useful, and therefore subject to be increased by natural selection.
Summary and Concluding Remarks on Hybridity.
The facts which are of the greatest importance to a comprehension of this very difficult subject are those which show the extreme susceptibility of the reproductive system both in plants and animals. We have seen how both these classes of organisms may be rendered infertile, by a change of conditions which does not affect their general health, by captivity, or by too close interbreeding. We have seen, also, that infertility is frequently correlated with a difference of colour, or with other characters; that it is not proportionate to divergence of structure; that it varies in reciprocal crosses between pairs of the same species; while in the cases of dimorphic and trimorphic plants the different crosses between the same pair of individuals may be fertile or sterile at the same time. It appears as if fertility depended on such a delicate adjustment of the male and female elements to each other, that, unless constantly kept up by the preservation of the most fertile individuals, sterility is always liable to arise. This preservation always occurs within the limits of each species, both because fertility is of the highest importance to the continuance of the race, and also because sterility (and to a less extent infertility) is self-destructive as well as injurious to the species.
So long therefore as a species remains undivided, and in occupation of a continuous area, its fertility is kept up by natural selection; but the moment it becomes separated, either by geographical or selective isolation, or by diversity of station or of habits, then, while each portion must be kept fertile inter se, there is nothing to prevent infertility arising between the two separated portions. As the two portions will necessarily exist under somewhat different conditions of life, and will usually have acquired some diversity of form and colour—both which circumstances we know to be either the cause of infertility or to be correlated with it,—the fact of some degree of infertility usually appearing between closely allied but locally or physiologically segregated species is exactly what we should expect.
The reason why varieties do not usually exhibit a similar amount of infertility is not difficult to explain. The popular conclusions on this matter have been drawn chiefly from what occurs among domestic animals, and we have seen that the very first essential to their becoming domesticated was that they should continue fertile under changed conditions of life. During the slow process of the formation of new varieties by conscious or unconscious selection, fertility has always been an essential character, and has thus been invariably preserved or increased; while there is some evidence to show that domestication itself tends to increase fertility.
Among plants, wild species and varieties have been more frequently experimented on than among animals, and we accordingly find numerous cases in which distinct species of plants are perfectly fertile when crossed, their hybrid offspring being also fertile inter se. We also find some few examples of the converse fact—varieties of the same species which when crossed are infertile or even sterile.
The idea that either infertility or geographical isolation is absolutely essential to the formation of new species, in order to prevent the swamping effects of intercrossing, has been shown to be unsound, because the varieties or incipient species will, in most cases, be sufficiently isolated by having adopted different habits or by frequenting different stations; while selective association, which is known to be general among distinct varieties or breeds of the same species, will produce an effective isolation even when the two forms occupy the same area.
From the various considerations now adverted to, Mr. Darwin arrived at the conclusion that the sterility or infertility of species with each other, whether manifested in the difficulty of obtaining first crosses between them or in the sterility of the hybrids thus obtained, is not a constant or necessary result of specific difference, but is incidental on unknown peculiarities of the reproductive system. These peculiarities constantly tend to arise under changed conditions owing to the extreme susceptibility of that system, and they are usually correlated with variations of form or of colour. Hence, as fixed differences of form and colour, slowly gained by natural selection in adaptation to changed conditions, are what essentially characterise distinct species, some amount of infertility between species is the usual result.
Here the problem was left by Mr. Darwin; but we have shown that its solution may be carried a step further. If we accept the association of some degree of infertility, however slight, as a not unfrequent accompaniment of the external differences which always arise in a state of nature between varieties and incipient species, it has been shown that natural selection has power to increase that infertility just as it has power to increase other favourable variations. Such an increase of infertility will be beneficial, whenever new species arise in the same area with the parent form; and we thus see how, out of the fluctuating and very unequal amounts of infertility correlated with physical variations, there may have arisen that larger and more constant amount which appears usually to characterise well-marked species.
The great body of facts of which a condensed account has been given in the present chapter, although from an experimental point of view very insufficient, all point to the general conclusion we have now reached, and afford us a not unsatisfactory solution of the great problem of hybridism in relation to the origin of species by means of natural selection. Further experimental research is needed in order to complete the elucidation of the subject; but until these additional facts are forthcoming no new theory seems required for the explanation of the phenomena.
FOOTNOTES:
[51] Darwin's Animals and Plants under Domestication, vol. ii. pp. 163-170.
[52] For a full account of these interesting facts and of the various problems to which they give rise, the reader must consult Darwin's volume on The Different Forms of Flowers in Plants of the same Species, chaps, i.-iv.
[53] See Nature, vol. xxi. p. 207.
[54] Low's Domesticated Animals of Great Britain, Introduction, p. lxiv.
[55] Low's Domesticated Animals, p. 28.
[56] Amaryllidaceae, by the Hon. and Rev. William Herbert, p. 379.
[57] Origin of Species, p. 239.
[58] Origin of Species, sixth edition, p. 9.
[59] In the Medico-Chirurgical Transactions, vol. liii. (1870), Dr. Ogle has adduced some curious physiological facts bearing on the presence or absence of white colours in the higher animals. He states that a dark pigment in the olfactory region of the nostrils is essential to perfect smell, and that this pigment is rarely deficient except when the whole animal is pure white, and the creature is then almost without smell or taste. He observes that there is no proof that, in any of the cases given above, the black animals actually eat the poisonous root or plant; and that the facts are readily understood if the senses of smell and taste are dependent on a pigment which is absent in the white animals, who therefore eat what those gifted with normal senses avoid. This explanation however hardly seems to cover the facts. We cannot suppose that almost all the sheep in the world (which are mostly white) are without smell or taste. The cutaneous disease on the white patches of hair on horses, the special liability of white terriers to distemper, of white chickens to the gapes, and of silkworms which produce yellow silk to the fungus, are not explained by it. The analogous facts in plants also indicate a real constitutional relation with colour, not an affection of the sense of smell and taste only.
[60] For all these facts, see Animals and Plants under Domestication, vol. ii. pp. 335-338.
[61] Animals and Plants under Domestication, vol. ii. pp. 102, 103.
[62] As this argument is a rather difficult one to follow, while its theoretical importance is very great, I add here the following briefer exposition of it, in a series of propositions; being, with a few verbal alterations, a copy of what I wrote on the subject about twenty years back. Some readers may find this easier to follow than the fuller discussion in the text:—
Can Sterility of Hybrids have been Produced by Natural Selection?
1. Let there be a species which has varied into two forms each adapted to certain existing conditions better than the parent form, which they soon supplant.
2. If these two forms, which are supposed to coexist in the same district, do not intercross, natural selection will accumulate all favourable variations till they become well suited to their conditions of life, and form two slightly differing species.
3. But if these two forms freely intercross with each other, and produce hybrids, which are also quite fertile inter se, then the formation of the two distinct races or species will be retarded, or perhaps entirely prevented; for the offspring of the crossed unions will be more vigorous owing to the cross, although less adapted to their conditions of life than either of the pure breeds.
4. Now, let a partial sterility of the hybrids of some considerable proportion of these two forms arise; and, as this would probably be due to some special conditions of life, we may fairly suppose it to arise in some definite portion of the area occupied by the two forms.
5. The result will be that, in that area, the hybrids (although continually produced by first crosses almost as freely as before) will not themselves increase so rapidly as the two pure forms; and as the two pure forms are, by the terms of the problem, better suited to their several conditions of life than the hybrids, they will inevitably increase more rapidly, and will continually tend to supplant the hybrids altogether at every recurrent severe struggle for existence.
6. We may fairly suppose, also, that as soon as any sterility appears some disinclination to cross unions will appear, and this will further tend to the diminution of the production of hybrids.
7. In the other part of the area, however, where hybridism occurs with perfect freedom, hybrids of various degrees may increase till they equal or even exceed in number the pure species—that is, the incipient species will be liable to be swamped by intercrossing.
8. The first result, then, of a partial sterility of crosses appearing in one part of the area occupied by the two forms, will be—that the great majority of the individuals will there consist of the two pure forms only, while in the remaining part these will be in a minority,—which is the same as saying that the new physiological variety of the two forms will be better suited to the conditions of existence than the remaining portion which has not varied physiologically.
9. But when the struggle for existence becomes severe, that variety which is best adapted to the conditions of existence always supplants that which is imperfectly adapted; therefore, by natural selection the varieties which are sterile when crossed will become established as the only ones.
10. Now let variations in the amount of sterilityand in the disinclination to crossed unions continue to occur—also in certain parts of the area: exactly the same result must recur, and the progeny of this new physiological variety will in time occupy the whole area.
11. There is yet another consideration that would facilitate the process. It seems probable that the sterility variations would, to some extent, concur with, and perhaps depend upon, the specific variations; so that, just in proportion as the two forms diverged and became better adapted to the conditions of existence, they would become more sterile when intercrossed. If this were the case, then natural selection would act with double strength; and those which were better adapted to survive both structurally and physiologically would certainly do so.
[63] Cases of this kind are referred to at p. 155. It must, however, be noted, that such sterility in first crosses appears to be equally rare between different species of the same genus as between individuals of the same species. Mules and other hybrids are freely produced between very distinct species, but are themselves infertile or quite sterile; and it is this infertility or sterility of the hybrids that is the characteristic—and was once thought to be the criterion—of species, not the sterility of their first crosses. Hence we should not expect to find any constant infertility in the first crosses between the distinct strains or varieties that formed the starting-point of new species, but only a slight amount of infertility in their mongrel offspring. It follows, that Mr. Romanes' theory of Physiological Selection—which assumes sterility or infertility between first crosses as the fundamental fact in the origin of species—does not accord with the general phenomena of hybridism in nature.
[64] The exact number is 1219.51, but the fractions are omitted for clearness.
CHAPTER VIII
THE ORIGIN AND USES OF COLOUR IN ANIMALS
The Darwinian theory threw new light on organic colour—The problem to be solved—The constancy of animal colour indicates utility—Colour and environment—Arctic animals white—Exceptions prove the rule—Desert, forest, nocturnal, and oceanic animals—General theories of animal colour—Variable protective colouring—Mr. Poulton's experiments—Special or local colour adaptations—Imitation of particular objects—How they have been produced—Special protective colouring of butterflies—Protective resemblance among marine animals—Protection by terrifying enemies—Alluring coloration—The coloration of birds' eggs—Colour as a means of recognition—Summary of the preceding exposition—Influence of locality or of climate on colour—Concluding remarks.
Among the numerous applications of the Darwinian theory in the interpretation of the complex phenomena presented by the organic world, none have been more successful, or are more interesting, than those which deal with the colours of animals and plants. To the older school of naturalists colour was a trivial character, eminently unstable and untrustworthy in the determination of species; and it appeared to have, in most cases, no use or meaning to the objects which displayed it. The bright and often gorgeous coloration of insect, bird, or flower, was either looked upon as having been created for the enjoyment of mankind, or as due to unknown and perhaps undiscoverable laws of nature.
But the researches of Mr. Darwin totally changed our point of view in this matter. He showed, clearly, that some of the colours of animals are useful, some hurtful to them; and he believed that many of the most brilliant colours were developed by sexual choice; while his great general principle, that all the fixed characters of organic beings have been developed under the action of the law of utility, led to the inevitable conclusion that so remarkable and conspicuous a character as colour, which so often constitutes the most obvious distinction of species from species, or group from group, must also have arisen from survival of the fittest, and must, therefore, in most cases have some relation to the wellbeing of its possessors. Continuous observation and research, carried on by multitudes of observers during the last thirty years, have shown this to be the case; but the problem is found to be far more complex than was at first supposed. The modes in which colour is of use to different classes of organisms is very varied, and have probably not yet been all discovered; while the infinite variety and marvellous beauty of some of its developments are such as to render it hopeless to arrive at a complete and satisfactory explanation of every individual case. So much, however, has been achieved, so many curious facts have been explained, and so much light has been thrown on some of the most obscure phenomena of nature, that the subject deserves a prominent place in any account of the Darwinian theory.
The Problem to be Solved.
Before dealing with the various modifications of colour in the animal world it is necessary to say a few words on colour in general, on its prevalence in nature, and how it is that the colours of animals and plants require any special explanation. What we term colour is a subjective phenomenon, due to the constitution of our mind and nervous system; while, objectively, it consists of light-vibrations of different wave-lengths emitted by, or reflected from, various objects. Every visible object must be coloured, because to be visible it must send rays of light to our eye. The kind of light it sends is modified by the molecular constitution or the surface texture of the object. Pigments absorb certain rays and reflect the remainder, and this reflected portion has to our eyes a definite colour, according to the portion of the rays constituting white light which are absorbed. Interference colours are produced either by thin films or by very fine striae on the surfaces of bodies, which cause rays of certain wave-lengths to neutralise each other, leaving the remainder to produce the effects of colour. Such are the colours of soap-bubbles, or of steel or glass on which extremely fine lines have been ruled; and these colours often produce the effect of metallic lustre, and are the cause of most of the metallic hues of birds and insects.
As colour thus depends on molecular or chemical constitution or on the minute surface texture of bodies, and, as the matter of which organic beings are composed consists of chemical compounds of great complexity and extreme instability, and is also subject to innumerable changes during growth and development, we might naturally expect the phenomena of colour to be more varied here than in less complex and more stable compounds. Yet even in the inorganic world we find abundant and varied colours; in the earth and in the water; in metals, gems, and minerals; in the sky and in the ocean; in sunset clouds and in the many-tinted rainbow. Here we can have no question of use to the coloured object, and almost as little perhaps in the vivid red of blood, in the brilliant colours of red snow and other low algae and fungi, or even in the universal mantle of green which clothes so large a portion of the earth's surface. The presence of some colour, or even of many brilliant colours, in animals and plants would require no other explanation than does that of the sky or the ocean, of the ruby or the emerald—that is, it would require a purely physical explanation only. It is the wonderful individuality of the colours of animals and plants that attracts our attention—the fact that the colours are localised in definite patterns, sometimes in accordance with structural characters, sometimes altogether independent of them; while often differing in the most striking and fantastic manner in allied species. We are thus compelled to look upon colour not merely as a physical but also as a biological characteristic, which has been differentiated and specialised by natural selection, and must, therefore, find its explanation in the principle of adaptation or utility.
The Constancy of Animal Colour indicates Utility.
That the colours and markings of animals have been acquired under the fundamental law of utility is indicated by a general fact which has received very little attention. As a rule, colour and marking are constant in each species of wild animal, while, in almost every domesticated animal, there arises great variability. We see this in our horses and cattle, our dogs and cats, our pigeons and poultry. Now, the essential difference between the conditions of life of domesticated and wild animals is, that the former are protected by man, while the latter have to protect themselves. The extreme variations in colour that immediately arise under domestication indicate a tendency to vary in this way, and the occasional occurrence of white or piebald or other exceptionally coloured individuals of many species in a state of nature, shows that this tendency exists there also; and, as these exceptionally coloured individuals rarely or never increase, there must be some constant power at work to keep it in check. This power can only be natural selection or the survival of the fittest, which again implies that some colours are useful, some injurious, in each particular case. With this principle as our guide, let us see how far we can account both for the general and special colours of the animal world.
Colour and Environment.
The fact that first strikes us in our examination of the colours of animals as a whole, is the close relation that exists between these colours and the general environment. Thus, white prevails among arctic animals; yellow or brown in desert species; while green is only a common colour in tropical evergreen forests. If we consider these cases somewhat carefully we shall find, that they afford us excellent materials for forming a judgment on the various theories that have been suggested to account for the colours of the animal world.
In the arctic regions there are a number of animals which are wholly white all the year round, or which only turn white in winter. Among the former are the polar bear and the American polar hare, the snowy owl and the Greenland falcon; among the latter the arctic fox, the arctic hare, the ermine, and the ptarmigan. Those which are permanently white remain among the snow nearly all the year round, while those which change their colour inhabit regions which are free from snow in summer. The obvious explanation of this style of coloration is, that it is protective, serving to conceal the herbivorous species from their enemies, and enabling carnivorous animals to approach their prey unperceived. Two other explanations have, however, been suggested. One is, that the prevalent white of the arctic regions has a direct effect in producing the white colour in animals, either by some photographic or chemical action on the skin or by a reflex action through vision. The other is, that the white colour is chiefly beneficial as a means of checking radiation and so preserving animal heat during the severity of an arctic winter. The first is part of the general theory that colour is the effect of coloured light on the objects—a pure hypothesis which has, I believe, no facts whatever to support it. The second suggestion is also an hypothesis merely, since it has not been proved by experiment that a white colour, per se, independently of the fur or feathers which is so coloured, has any effect whatever in checking the radiation of low-grade heat like that of the animal body. But both alike are sufficiently disproved by the interesting exceptions to the rule of white coloration in the arctic regions, which exceptions are, nevertheless, quite in harmony with the theory of protection.
Whenever we find arctic animals which, from whatever cause, do not require protection by the white colour, then neither the cold nor the snow-glare has any effect upon their coloration. The sable retains its rich brown fur throughout the Siberian winter; but it frequents trees at that season and not only feeds partially on fruits or seeds, but is able to catch birds among the branches of the fir-trees, with the bark of which its colour assimilates. Then we have that thoroughly arctic animal, the musk-sheep, which is brown and conspicuous; but this animal is gregarious, and its safety depends on its association in small herds. It is, therefore, of more importance for it to be able to recognise its kind at a distance than to be concealed from its enemies, against which it can well protect itself so long as it keeps together in a compact body. But the most striking example is that of the common raven, which is a true arctic bird, and is found even in mid-winter as far north as any known bird or mammal. Yet it always retains its black coat, and the reason, from our point of view, is obvious. The raven is a powerful bird and fears no enemy, while, being a carrion-feeder, it has no need for concealment in order to approach its prey. The colour of the raven and of the musk-sheep are, therefore, both inconsistent with any other theory than that the white colour of arctic animals has been acquired for concealment, and to that theory both afford a strong support. Here we have a striking example of the exception proving the rule.
In the desert regions of the earth we find an even more general accordance of colour with surroundings. The lion, the camel, and all the desert antelopes have more or less the colour of the sand or rock among which they live. The Egyptian cat and the Pampas cat are sandy or earth coloured. The Australian kangaroos are of similar tints, and the original colour of the wild horse is supposed to have been sandy or clay coloured. Birds are equally well protected by assimilative hues; the larks, quails, goatsuckers, and grouse which abound in the North African and Asiatic deserts are all tinted or mottled so as closely to resemble the average colour of the soil in the districts they inhabit. Canon Tristram, who knows these regions and their natural history so well, says, in an often quoted passage: "In the desert, where neither trees, brushwood, nor even undulations of the surface afford the slightest protection to its foes, a modification of colour which shall be assimilated to that of the surrounding country is absolutely necessary. Hence, without exception, the upper plumage of every bird, whether lark, chat, sylvain, or sand-grouse, and also the fur of all the smaller mammals, and the skin of all the snakes and lizards, is of one uniform isabelline or sand colour."
Passing on to the tropical regions, it is among their evergreen forests alone that we find whole groups of birds whose ground colour is green. Parrots are very generally green, and in the East we have an extensive group of green fruit-eating pigeons; while the barbets, bee-eaters, turacos, leaf-thrushes (Phyllornis), white-eyes (Zosterops), and many other groups, have so much green in their plumage as to tend greatly to their concealment among the dense foliage. There can be no doubt that these colours have been acquired as a protection, when we see that in all the temperate regions, where the leaves are deciduous, the ground colour of the great majority of birds, especially on the upper surface, is a rusty brown of various shades, well corresponding with the bark, withered leaves, ferns, and bare thickets among which they live in autumn and winter, and especially in early spring when so many of them build their nests.
Nocturnal animals supply another illustration of the same rule, in the dusky colours of mice, rats, bats, and moles, and in the soft mottled plumage of owls and goatsuckers which, while almost equally inconspicuous in the twilight, are such as to favour their concealment in the daytime.
An additional illustration of general assimilation of colour to the surroundings of animals, is furnished by the inhabitants of the deep oceans. Professor Moseley of the Challenger Expedition, in his British Association lecture on this subject, says: "Most characteristic of pelagic animals is the almost crystalline transparency of their bodies. So perfect is this transparency that very many of them are rendered almost entirely invisible when floating in the water, while some, even when caught and held up in a glass globe, are hardly to be seen. The skin, nerves, muscles, and other organs are absolutely hyaline and transparent, but the liver and digestive tract often remain opaque and of a yellow or brown colour, and exactly resemble when seen in the water small pieces of floating seaweed." Such marine organisms, however, as are of larger size, and either occasionally or habitually float on the surface, are beautifully tinged with blue above, thus harmonising with the colour of the sea as seen by hovering birds; while they are white below, and are thus invisible against the wave-foam and clouds as seen by enemies beneath the surface. Such are the tints of the beautiful nudibranchiate mollusc, Glaucus atlanticus, and many others.
General Theories of Animal Colour.
We are now in a position to test the general theories, or, to speak more correctly, the popular notions, as to the origin of animal coloration, before proceeding to apply the principle of utility to the explanation of some among the many extraordinary manifestations of colour in the animal world. The most generally received theory undoubtedly is, that brilliancy and variety of colour are due to the direct action of light and heat; a theory no doubt derived from the abundance of bright-coloured birds, insects, and flowers which are brought from tropical regions. There are, however, two strong arguments against this theory. We have already seen how generally bright coloration is wanting in desert animals, yet here heat and light are both at a maximum, and if these alone were the agents in the production of colour, desert animals should be the most brilliant. Again, all naturalists who have lived in tropical regions know that the proportion of bright to dull coloured species is little if any greater there than in the temperate zone, while there are many tropical groups in which bright colours are almost entirely unknown. No part of the world presents so many brilliant birds as South America, yet there are extensive families, containing many hundreds of species, which are as plainly coloured as our average temperate birds. Such are the families of the bush-shrikes and ant-thrushes (Formicariidae), the tyrant-shrikes (Tyrannidae), the American creepers (Dendrocolaptidae), together with a large proportion of the wood-warblers (Mniotiltidae), the finches, the wrens, and some other groups. In the eastern hemisphere, also, we have the babbling-thrushes (Timaliidae), the cuckoo-shrikes (Campephagidae), the honey-suckers (Meliphagidae), and several other smaller groups which are certainly not coloured above the average standard of temperate birds.
Again, there are many families of birds which spread over the whole world, temperate and tropical, and among these the tropical species rarely present any exceptional brilliancy of colour. Such are the thrushes, goatsuckers, hawks, plovers, and ducks; and in the last-named group it is the temperate and arctic zones that afford the most brilliant coloration.
The same general facts are found to prevail among insects. Although tropical insects present some of the most gorgeous coloration in the whole realm of nature, yet there are thousands and tens of thousands of species which are as dull coloured as any in our cloudy land. The extensive family of the carnivorous ground-beetles (Carabidae) attains its greatest brilliancy in the temperate zone; while by far the larger proportion of the great families of the longicorns and the weevils, are of obscure colours even in the tropics. In butterflies, there is undoubtedly a larger proportion of brilliant colour in the tropics; but if we compare families which are almost equally developed over the globe—as the Pieridae or whites and yellows, and the Satyridae or ringlets—we shall find no great disproportion in colour between those of temperate and tropical regions.
The various facts which have now briefly been noticed are sufficient to indicate that the light and heat of the sun are not the direct causes of the colours of animals, although they may favour the production of colour when, as in tropical regions, the persistent high temperature favours the development of the maximum of life. We will now consider the next suggestion, that light reflected from surrounding coloured objects tends to produce corresponding colours in the animal world.
This theory is founded on a number of very curious facts which prove, that such a change does sometimes occur and is directly dependent on the colours of surrounding objects; but these facts are comparatively rare and exceptional in their nature, and the same theory will certainly not apply to the infinitely varied colours of the higher animals, many of which are exposed to a constantly varying amount of light and colour during their active existence. A brief sketch of these dependent changes of colour may, however, be advantageously given here.
Variable Protective Colouring.
There are two distinct kinds of change of colour in animals due to the colouring of the environment. In one case the change is caused by reflex action set up by the animal seeing the colour to be imitated, and the change produced can be altered or repeated as the animal changes its position. In the other case the change occurs but once, and is probably not due to any conscious or sense action, but to some direct influence on the surface tissues while the creature is undergoing a moult or change to the pupa form.
The most striking example of the first class is that of the chameleon, which changes to white, brown, yellowish, or green, according to the colour of the object on which it rests. This change is brought about by means of two layers of pigment cells, deeply seated in the skin, and of bluish and yellowish colours. By suitable muscles these cells can be forced upwards so as to modify the colour of the skin, which, when they are not brought into action, is a dirty white. These animals are excessively sluggish and defenceless, and the power of changing their colour to that of their immediate surroundings is no doubt of great service to them. Many of the flatfish are also capable of changing their colour according to the colour of the bottom they rest on; and frogs have a similar power to a limited extent. Some crustacea also change colour, and the power is much developed in the Chameleon shrimp (Mysis Chamaeleon) which is gray when on sand, but brown or green when among brown or green seaweed. It has been proved by experiment that when this animal is blinded the change does not occur. In all these cases, therefore, we have some form of reflex or sense action by which the change is produced, probably by means of pigment cells beneath the skin as in the chameleon.
The second class consists of certain larvae, and pupae, which undergo changes of colour when exposed to differently coloured surroundings. This subject has been carefully investigated by Mr. E.B. Poulton, who has communicated the results of his experiments to the Royal Society.[65] It had been noticed that some species of larvae which fed on several different plants had colours more or less corresponding to the particular plant the individual fed on. Numerous cases are given in Professor Meldola's article on "Variable Protective Colouring" (Proc. Zool. Soc., 1873, p. 153), and while the general green coloration was attributed to the presence of chlorophyll beneath the skin, the particular change in correspondence to each food-plant was attributed to a special function which had been developed by natural selection. Later on, in a note to his translation of Weissmann's Theory of Descent, Professor Meldola seemed disposed to think that the variations of colour of some of the species might be phytophagic—that is, due to the direct action of the differently coloured leaves on which the insect fed. Mr. Poulton's experiments have thrown much light on this question, since he has conclusively proved that, in the case of the sphinx caterpillar of Smerinthus ocellatus, the change of colour is not due to the food but to the coloured light reflected from the leaves.
This was shown by feeding two sets of larvae on the same plant but exposed to differently coloured surroundings, obtained by sewing the leaves together, so that in one case only the dark upper surface, in the other the whitish under surface was exposed to view. The result in each case was a corresponding change of colour in the larvae, confirming the experiments on different individuals of the same batch of larvae which had been supplied with different food-plants or exposed to a different coloured light.
An even more interesting series of experiments was made on the colours of pupae, which in many cases were known to be affected by the material on which they underwent their transformations. The late Mr. T.W. Wood proved, in 1867, that the pupae of the common cabbage butterflies (Pieris brassicae and P. rapae) were either light, or dark, or green, according to the coloured boxes they were kept in, or the colours of the fences, walls, etc., against which they were suspended. Mrs. Barber in South Africa found that the pupae of Papilio Nireus underwent a similar change, being deep green when attached to orange leaves of the same tint, pale yellowish-green when on a branch of the bottle-brush tree whose half-dried leaves were of this colour, and yellowish when attached to the wooden frame of a box. A few other observers noted similar phenomena, but nothing more was done till Mr. Poulton's elaborate series of experiments with the larvae of several of our common butterflies were the means of clearing up several important points. He showed that the action of the coloured light did not affect the pupa itself but the larva, and that only for a limited period of time. After a caterpillar has done feeding it wanders about seeking a suitable place to undergo its transformation. When this is found it rests quietly for a day or two, spinning the web from which it is to suspend itself; and it is during this period of quiescence, and perhaps also the first hour or two after its suspension, that the action of the surrounding coloured surfaces determines, to a considerable extent, the colour of the pupa. By the application of various surrounding colours during this period, Mr. Poulton was able to modify the colour of the pupa of the common tortoise-shell butterfly from nearly black to pale, or to a brilliant golden; and that of Pieris rapae from dusky through pinkish to pale green. It is interesting to note, that the colours produced were in all cases such only as assimilated with the surroundings usually occupied by the species, and also, that colours which did not occur in such surroundings, as dark red or blue, only produced the same effects as dusky or black.
Careful experiments were made to ascertain whether the effect was produced through the sight of the caterpillar. The ocelli were covered with black varnish, but neither this, nor cutting off the spines of the tortoise-shell larva to ascertain whether they might be sense-organs, produced any effect on the resulting colour. Mr. Poulton concludes, therefore, that the colour-action probably occurs over the whole surface of the body, setting up physiological processes which result in the corresponding colour-change of the pupa. Such changes are, however, by no means universal, or even common, in protectively coloured pupae, since in Papilio machaon and some others which have been experimented on, both in this country and abroad, no change can be produced on the pupa by any amount of exposure to differently coloured surroundings. It is a curious point that, with the small tortoise-shell larva, exposure to light from gilded surfaces produced pupae with a brilliant golden lustre; and the explanation is supposed to be that mica abounded in the original habitat of the species, and that the pupae thus obtained protection when suspended against micaceous rock. Looking, however, at the wide range of the species and the comparatively limited area in which micaceous rocks occur, this seems a rather improbable explanation, and the occurrence of this metallic appearance is still a difficulty. It does not, however, commonly occur in this country in a natural state.
The two classes of variable colouring here discussed are evidently exceptional, and can have little if any relation to the colours of those more active creatures which are continually changing their position with regard to surrounding objects, and whose colours and markings are nearly constant throughout the life of the individual, and (with the exception of sexual differences) in all the individuals of the species. We will now briefly pass in review the various characteristics and uses of the colours which more generally prevail in nature; and having already discussed those protective colours which serve to harmonise animals with their general environment, we have to consider only those cases in which the colour resemblance is more local or special in its character.
Special or Local Colour Adaptations.
This form of colour adaptation is generally manifested by markings rather than by colour alone, and is extremely prevalent both among insects and vertebrates, so that we shall be able to notice only a few illustrative cases. Among our native birds we have the snipe and woodcock, whose markings and tints strikingly accord with the dead marsh vegetation among which they live; the ptarmigan in its summer dress is mottled and tinted exactly like the lichens which cover the stones of the higher mountains; while young unfledged plovers are spotted so as exactly to resemble the beach pebbles among which they crouch for protection, as beautifully exhibited in one of the cases of British birds in the Natural History Museum at South Kensington.
In mammalia, we notice the frequency of rounded spots on forest or tree haunting animals of large size, as the forest deer and the forest cats; while those that frequent reedy or grassy places are striped vertically, as the marsh antelopes and the tiger. I had long been of opinion that the brilliant yellow and black stripes of the tiger were adaptive, but have only recently obtained proof that it is so. An experienced tiger-hunter, Major Walford, states in a letter, that the haunts of the tiger are invariably full of the long grass, dry and pale yellow for at least nine months of the year, which covers the ground wherever there is water in the rainy season, and he adds: "I once, while following up a wounded tiger, failed for at least a minute to see him under a tree in grass at a distance of about twenty yards—jungle open—but the natives saw him, and I eventually made him out well enough to shoot him, but even then I could not see at what part of him I was aiming. There can be no doubt whatever that the colour of both the tiger and the panther renders them almost invisible, especially in a strong blaze of light, when among grass, and one does not seem to notice stripes or spots till they are dead." It is the black shadows of the vegetation that assimilate with the black stripes of the tiger; and, in like manner, the spotty shadows of leaves in the forest so harmonise with the spots of ocelots, jaguars, tiger-cats, and spotted deer as to afford them a very perfect concealment.
In some cases the concealment is effected by colours and markings which are so striking and peculiar that no one who had not seen the creature in its native haunts would imagine them to be protective. An example of this is afforded by the banded fruit pigeon of Timor, whose pure white head and neck, black wings and back, yellow belly, and deeply-curved black band across the breast, render it a very handsome and conspicuous bird. Yet this is what Mr. H.O. Forbes says of it: "On the trees the white-headed fruit pigeon (Ptilopus cinctus) sate motionless during the heat of the day in numbers, on well-exposed branches; but it was with the utmost difficulty that I or my sharp-eyed native servant could ever detect them, even in trees where we knew they were sitting."[66] The trees referred to are species of Eucalyptus which abound in Timor. They have whitish or yellowish bark and very open foliage, and it is the intense sunlight casting black curved shadows of one branch upon another, with the white and yellow bark and deep blue sky seen through openings of the foliage, that produces the peculiar combination of colours and shadows to which the colours and markings of this bird have become so closely assimilated.
Even such brilliant and gorgeously coloured birds as the sun-birds of Africa are, according to an excellent observer, often protectively coloured. Mrs. M.E. Barber remarks that "A casual observer would scarcely imagine that the highly varnished and magnificently coloured plumage of the various species of Noctarinea could be of service to them, yet this is undoubtedly the case. The most unguarded moments of the lives of these birds are those that are spent amongst the flowers, and it is then that they are less wary than at any other time. The different species of aloes, which blossom in succession, form the principal sources of their winter supplies of food; and a legion of other gay flowering plants in spring and summer, the aloe blossoms especially, are all brilliantly coloured, and they harmonise admirably with the gay plumage of the different species of sun-birds. Even the keen eye of a hawk will fail to detect them, so closely do they resemble the flowers they frequent. The sun-birds are fully aware of this fact, for no sooner have they relinquished the flowers than they become exceedingly wary and rapid in flight, darting arrow-like through the air and seldom remaining in exposed situations. The black sun-bird (Nectarinea amethystina) is never absent from that magnificent forest-tree, the 'Kaffir Boom' (Erythrina caffra); all day long the cheerful notes of these birds may be heard amongst its spreading branches, yet the general aspect of the tree, which consists of a huge mass of scarlet and purple-black blossoms without a single green leaf, blends and harmonises with the colours of the black sun-bird to such an extent that a dozen of them may be feeding amongst its blossoms without being conspicuous, or even visible."[67]
Some other cases will still further illustrate how the colours of even very conspicuous animals may be adapted to their peculiar haunts.
The late Mr. Swinhoe says of the Kerivoula picta, which he observed in Formosa: "The body of this bat was of an orange colour, but the wings were painted with orange-yellow and black. It was caught suspended, head downwards, on a cluster of the fruit of the longan tree (Nephelium longanum). Now this tree is an evergreen, and all the year round some portion of its foliage is undergoing decay, the particular leaves being, in such a stage, partially orange and black. This bat can, therefore, at all seasons suspend from its branches and elude its enemies by its resemblance to the leaves of the tree."[68]
Even more curious is the case of the sloths—defenceless animals which feed upon leaves, and hang from the branches of trees with their back downwards. Most of the species have a curious buff-coloured spot on the back, rounded or oval in shape and often with a darker border, which seems placed there on purpose to make them conspicuous; and this was a great puzzle to naturalists, because the long coarse gray or greenish hair was evidently like tree-moss and therefore protective. But an old writer, Baron von Slack, in his Voyage to Surinam (1810), had already explained the matter. He says: "The colour and even the shape of the hair are much like withered moss, and serve to hide the animal in the trees, but particularly when it has that orange-coloured spot between the shoulders and lies close to the tree; it looks then exactly like a piece of branch where the rest has been broken off, by which the hunters are often deceived." Even such a huge animal as the giraffe is said to be perfectly concealed by its colour and form when standing among the dead and broken trees that so often occur on the outskirts of the thickets where it feeds. The large blotch-like spots on the skin and the strange shape of the head and horns, like broken branches, so tend to its concealment that even the keen-eyed natives have been known to mistake trees for giraffes or giraffes for trees.
Innumerable examples of this kind of protective colouring occur among insects; beetles mottled like the bark of trees or resembling the sand or rock or moss on which they live, with green caterpillars of the exact general tints of the foliage they feed on; but there are also many cases of detailed imitation of particular objects by insects that must be briefly described.[69]
Protective Imitation of Particular Objects.
The insects which present this kind of imitation most perfectly are the Phasmidae, or stick and leaf insects. The well-known leaf-insects of Ceylon and of Java, species of Phyllium, are so wonderfully coloured and veined, with leafy expansions on the legs and thorax, that not one person in ten can see them when resting on the food-plant close beneath their eyes. Others resemble pieces of stick with all the minutiae of knots and branches, formed by the insects' legs, which are stuck out rigidly and unsymmetrically. I have often been unable to distinguish between one of these insects and a real piece of stick, till I satisfied myself by touching it and found it to be alive. One species, which was brought me in Borneo, was covered with delicate semitransparent green foliations, exactly resembling the hepaticae which cover pieces of rotten stick in the damp forests. Others resemble dead leaves in all their varieties of colour and form; and to show how perfect is the protection obtained and how important it is to the possessors of it, the following incident, observed by Mr. Belt in Nicaragua, is most instructive. Describing the armies of foraging ants in the forest which devour every insect they can catch, he says: "I was much surprised with the behaviour of a green leaf-like locust. This insect stood immovably among a host of ants, many of which ran over its legs without ever discovering there was food within their reach. So fixed was its instinctive knowledge that its safety depended on its immovability, that it allowed me to pick it up and replace it among the ants without making a single effort to escape. This species closely resembles a green leaf."[70]
Caterpillars also exhibit a considerable amount of detailed resemblance to the plants on which they live. Grass-feeders are striped longitudinally, while those on ordinary leaves are always striped obliquely. Some very beautiful protective resemblances are shown among the caterpillars figured in Smith and Abbott's Lepidopterous Insects of Georgia, a work published in the early part of the century, before any theories of protection were started. The plates in this work are most beautifully executed from drawings made by Mr. Abbott, representing the insects, in every case, on the plants which they frequented, and no reference is made in the descriptions to the remarkable protective details which appear upon the plates. We have, first, the larva of Sphinx fuciformis feeding on a plant with linear grass-like leaves and small blue flowers; and we find the insect of the same green as the leaves, striped longitudinally in accordance with the linear leaves, and with the head blue corresponding both in size and colour with the flowers. Another species (Sphinx tersa) is represented feeding on a plant with small red flowers situated in the axils of the leaves; and the larva has a row of seven red spots, unequal in size, and corresponding very closely with the colour and size of the flowers. Two other figures of sphinx larvae are very curious. That of Sphinx pampinatrix feeds on a wild vine (Vitis indivisa), having green tendrils, and in this species the curved horn on the tail is green, and closely imitates in its curve the tip of the tendril. But in another species (Sphinx cranta), which feeds on the fox-grape (Vitis vulpina), the horn is very long and red, corresponding with the long red-tipped tendrils of the plant. Both these larvae are green with oblique stripes, to harmonise with the veined leaves of the vines; but a figure is also given of the last-named species after it has done feeding, when it is of a decided brown colour and has entirely lost its horn. This is because it then descends to the ground to bury itself, and the green colour and red horn would be conspicuous and dangerous; it therefore loses both at the last moult. Such a change of colour occurs in many species of caterpillars. Sometimes the change is seasonal; and, in those which hibernate with us, the colour of some species, which is brownish in autumn in adaptation to the fading foliage, becomes green in spring to harmonise with the newly-opened leaves at that season.[71]
Some of the most curious examples of minute imitation are afforded by the caterpillars of the geometer moths, which are always brown or reddish, and resemble in form little twigs of the plant on which they feed. They have the habit, when at rest, of standing out obliquely from the branch, to which they hold on by their hind pair of prolegs or claspers, and remain motionless for hours. Speaking of these protective resemblances Mr. Jenner Weir says: "After being thirty years an entomologist I was deceived myself, and took out my pruning scissors to cut from a plum tree a spur which I thought I had overlooked. This turned out to be the larva of a geometer two inches long. I showed it to several members of my family, and defined a space of four inches in which it was to be seen, but none of them could perceive that it was a caterpillar."[72]
One more example of a protected caterpillar must be given. Mr. A. Everett, writing from Sarawak, Borneo, says: "I had a caterpillar brought me, which, being mixed by my boy with some other things, I took to be a bit of moss with two exquisite pinky-white seed-capsules; but I soon saw that it moved, and examining it more closely found out its real character: it is covered with hair, with two little pink spots on the upper surface, the general hue being more green. Its motions are very slow, and when eating the head is withdrawn beneath a fleshy mobile hood, so that the action of feeding does not produce any movement externally. It was found in the limestone hills at Busan, the situation of all others where mosses are most plentiful and delicate, and where they partially clothe most of the protruding masses of rock."
How these Imitations have been Produced.
To many persons it will seem impossible that such beautiful and detailed resemblances as those now described—and these are only samples of thousands that occur in all parts of the world—can have been brought about by the preservation of accidental useful variations. But this will not seem so surprising if we keep in mind the facts set forth in our earlier chapters—the rapid multiplication, the severe struggle for existence, and the constant variability of these and all other organisms. And, further, we must remember that these delicate adjustments are the result of a process which has been going on for millions of years, and that we now see the small percentage of successes among the myriads of failures. From the very first appearance of insects and their various kinds of enemies the need of protection arose, and was usually most easily met by modifications of colour. Hence, we may be sure that the earliest leaf-eating insects acquired a green colour as one of the necessities of their existence; and, as the species became modified and specialised, those feeding on particular species of plants would rapidly acquire the peculiar tints and markings best adapted to conceal them upon those plants. Then, every little variation that, once in a hundred years perhaps, led to the preservation of some larva which was thereby rather better concealed than its fellows, would form the starting-point of a further development, leading ultimately to that perfection of imitation in details which now astonishes us. The researches of Dr. Weismann illustrate this progressive adaptation. The very young larvae of several species are green or yellowish without any markings; they then, in subsequent moults, obtain certain markings, some of which are often lost again before the larva is fully grown. The early stages of those species which, like elephant hawk-moths (Chaerocampa), have the anterior segments elongated and retractile, with large eye-like spots to imitate the head of a vertebrate, are at first like those of non-retractile species, the anterior segments being as large as the rest. After the first moult they become smaller, comparatively; but it is only after the second moult that the ocelli begin to appear, and these are not fully defined till after the third moult. This progressive development of the individual—the ontogeny—gives us a clue to the ancestral development of the whole race—the phylogeny; and we are enabled to picture to ourselves the very slow and gradual steps by which the existing perfect adaptation has been brought about. In many larvae great variability still exists, and in some there are two or more distinctly-coloured forms—usually a dark and a light or a brown and a green form. The larva of the humming-bird hawk-moth (Macroglossa stellatarum) varies in this manner, and Dr. Weismann raised five varieties from a batch of eggs from one moth. It feeds on species of bedstraw (Galium verum and G. mollugo), and as the green forms are less abundant than the brown, it has probably undergone some recent change of food-plant or of habits which renders brown the more protective colour.
Special Protective Colouring of Butterflies.
We will now consider a few cases of special protective colouring in the perfect butterfly or moth. Mr. Mansel Weale states that in South Africa there is a great prevalence of white and silvery foliage or bark, sometimes of dazzling brilliancy, and that many insects and their larvae have brilliant silvery tints which are protective, among them being three species of butterflies whose undersides are silvery, and which are thus effectually protected when at rest.[73] A common African butterfly (Aterica meleagris) always settles on the ground with closed wings, which so closely resemble the soil of the district that it can with difficulty be seen, and the colour varies with the soil in different localities. Thus specimens from Senegambia were dull brown, the soil being reddish sand and iron-clay; those from Calabar and Cameroons were light brown with numerous small white spots, the soil of those countries being light brown clay with small quartz pebbles; while in other localities where the colours of the soil were more varied the colours of the butterfly varied also. Here we have variation in a single species which has become specialised in certain areas to harmonise with the colour of the soil.[74]
Many butterflies, in all parts of the world, resemble dead leaves on their under side, but those in which this form of protection is carried to the greatest perfection are the species of the Eastern genus Kallima. In India K. inachis, and in the larger Malay islands K. paralekta, are very common. They are rather large and showy butterflies, orange and bluish on the upper side, with a very rapid flight, and frequenting dry forests. Their habit is to settle always where there is some dead or decaying foliage, and the shape and colour of the wings (on the under surface), together with the attitude of the insect, is such as to produce an absolutely perfect imitation of a dead leaf. This is effected by the butterfly always settling on a twig, with the short tail of the hind wings just touching it and forming the leaf-stalk. From this a dark curved line runs across to the elongated tip of the upper wings, imitating the midrib, on both sides of which are oblique lines, formed partly by the nervures and partly by markings, which give the effect of the usual veining of a leaf. The head and antennae fit exactly between the closed upper wings so as not to interfere with the outline, which has just that amount of irregular curvature that is seen in dry and withered leaves. The colour is very remarkable for its extreme amount of variability, from deep reddish-brown to olive or pale yellow, hardly two specimens being exactly alike, but all coming within the range of colour of leaves in various stages of decay. Still more curious is the fact that the paler wings, which imitate leaves most decayed, are usually covered with small black dots, often gathered into circular groups, and so exactly resembling the minute fungi on decaying leaves that it is hard at first to believe that the insects themselves are not attacked by some such fungus. The concealment produced by this wonderful imitation is most complete, and in Sumatra I have often seen one enter a bush and then disappear like magic. Once I was so fortunate as to see the exact spot on which the insect settled; but even then I lost sight of it for some time, and only after a persistent search discovered that it was close before my eyes.[75] Here we have a kind of imitation, which is very common in a less developed form, carried to extreme perfection, with the result that the species is very abundant over a considerable area of country.
Protective Resemblance among Marine Animals.
Among marine animals this form of protection is very common. Professor Moseley tells us that all the inhabitants of the Gulf-weed are most remarkably coloured, for purposes of protection and concealment, exactly like the weed itself. "The shrimps and crabs which swarm in the weed are of exactly the same shade of yellow as the weed, and have white markings upon their bodies to represent the patches of Membranipora. The small fish, Antennarius, is in the same way weed-colour with white spots. Even a Planarian worm, which lives in the weed, is similarly yellow-coloured, and also a mollusc, Scyllaea pelagica." The same writer tells us that "a number of little crabs found clinging to the floats of the blue-shelled mollusc, Ianthina, were all coloured of a corresponding blue for concealment."[76]
Professor E.S. Morse of Salem, Mass., found that most of the New England marine mollusca were protectively coloured; instancing among others a little red chiton on rocks clothed with red calcareous algae, and Crepidula plana, living within the apertures of the shells of larger species of Gasteropods and of a pure white colour corresponding to its habitat, while allied species living on seaweed or on the outside of dark shells were dark brown.[77] A still more interesting case has been recorded by Mr. George Brady. He says: "Amongst the Nullipore which matted together the laminaria roots in the Firth of Clyde were living numerous small starfishes (Ophiocoma bellis) which, except when their writhing movements betrayed them, were quite undistinguishable from the calcareous branches of the alga; their rigid angularly twisted rays had all the appearance of the coralline, and exactly assimilated to its dark purple colour, so that though I held in my hand a root in which were half a dozen of the starfishes, I was really unable to detect them until revealed by their movements."[78]
These few examples are sufficient to show that the principle of protective coloration extends to the ocean as well as over the earth; and if we consider how completely ignorant we are of the habits and surroundings of most marine animals, it may well happen that many of the colours of tropical fishes, which seem to us so strange and so conspicuous, are really protective, owing to the number of equally strange and brilliant forms of corals, sea-anemones, sponges, and seaweeds among which they live.