Part 9

Another group, which agrees in colour with the general surroundings, is that of the 'glass-animals,' as they have been called, though perhaps 'crystal animals' is a better term. A great number of simple free-swimming marine forms, and a few fresh-water ones, are quite colourless, and perfectly transparent, or have at most a bluish or greenish tinge, and on this account they are quite invisible as long as they remain in the water. In our lakes there lives a little crustacean about a centimetre in length, of the order of water-fleas (_Leptodora hyalina_), a mighty hunter among the smallest animals, which swims forward jerkily with its long swimming-appendages, and widely spreads its six pairs of claws, armed with thorny bristles, like a weir basket, to seize its prey. We may have dozens of these in a glass of water without being able to see a single one, even when we hold the glass against the light, for the creatures are crystal-clear and transparent, and have exactly the same refractive power as the water. It requires a very sharp scrutiny and a knowledge of the animals to be able to detect in the water little yellowish stripes, which are the stomachs of the animals filled with food in process of digestion, for which, as we can readily understand, invisibility cannot very well be arranged. If the water be then strained through a fine cloth, a little gelatine-like mass of the bodies of the _Leptodora_ will remain on the sieve.

A great many of the lower marine animals are equally transparent, and as clear as water; most of the lower Medusæ, the ctenophores, various molluscs, the barrel-shaped Salpæ, worms, many crustaceans of quite different orders, and above all an enormous number of larvæ of the most diverse animal groups. I can remember seeing the sea at the shore at Mentone so full of Salpæ, that in every glass of sea-water drawn at random there were many of them, and sometimes a glass held a positive animal soup. But one did not see them in the glass of water, and only those who knew what to look for recognized them by the bluish intestinal sac that lies posteriorly in the invisible body. But when the water was poured off through a fine net, there remained on the filter a large mass of a crystalline gelatinous substance.

It is obvious that this must serve as a protective arrangement, for the animals are not seen by their pursuers; but it is not an _absolute_ protection, for they have many pursuers who do not wait till they see their prey, but are almost constantly snapping the mouth open and shut, leaving it to chance to bring them their prey. _No protective arrangement, however, affords absolute security_; it protects against some enemies, perhaps against many, but never against all.

But now let us turn to a group of a different colouring, the green animals. We are familiar with our big grass-green grasshopper, and we know how easily it is overlooked when it sits quietly on a high grass-stem, surrounded by grasses and herbage; the light grass-green of its whole body protects it most effectively from discovery: for myself, at least, I must confess that in a flowery meadow I have stood right in front of one, and have looked close to it for a long time without detecting it. In the same way countless insects of the most diverse groups--bugs, dipterous flies, sawflies, butterflies--and especially the larvæ (caterpillars) of the last, are of the same green as the plants on which they live, and this again applies to the predaceous species, as well as the species preyed upon. Thus the rapacious praying-mantis (_Mantis religiosa_) is as green as the grass in which it lurks motionless for its victim--a dragonfly, a fly, or a butterfly.

There are also green spiders, green amphibians like the edible frog, and especially the tree-frog, green reptiles like lizards and the tree-snakes of tropical forests. It is always animals which live among green that are green in colour.

We may wonder, for a moment, why there are so few green birds, since they spend so much of their time among the green leaves. But this paucity of green birds is only true of temperate climates. In Germany we have only the green woodpecker, the siskin, and a few other little birds, and even these are not of a bright green, but are rather greyish-green. The explanation lies in the long winter, when the trees are leafless. In the evergreen forests of the tropics there are numerous green birds belonging to very diverse families.

Yet another group with a common colour-adaptation deserves mention--the beasts of the night. They are all more or less grey, brown, yellowish, or a mixture of these colours, and it is obvious that, in the duskiness of night, they must blend better with their environment on this account. White mice and white rats cannot exist under natural conditions, since they are conspicuous in the night, and the same would be true of white bats, nightjars, and owls; but all of these have a coloration suited to nocturnal habits.

A very remarkable fact is that in many animals the colour-adaptation is a double one. Thus the Arctic fox is white only in winter, while in summer he is greyish-brown; the ermine changes in the same way, and the great white snowy owl of the Arctic regions has in summer a grey-brown variegated plumage. Many animals which are subject to persecution also change colour with the seasons, like the mountain hare (_Lepus variabilis_), which is brown in summer and pure white in winter, the Lapland lemming, and the ptarmigan (_Lagopus alpinus_), which do the same. It has been doubted whether natural selection can explain this double coloration, but I do not know where the difficulty lies, and there is certainly no other principle whose aid we can evoke. The mountain hare must have had some sort of colour before it attained to seasonal dimorphism. Let us assume that it was brown, that the climate became colder and the winter longer, then those hares would have most chance of surviving which became lighter in winter, and so a white race was formed. Poulton has shown that the whiteness is due to the fact that the dark hairs of the summer coat grow white as they lengthen at the beginning of winter, and the abundance of new hairs which complete the winter coat are from the first white throughout. If the white hairs were to persist throughout the summer it would be very disadvantageous to their wearer; so a double selection must take place, in summer the individuals which remain white, in winter those which remain brown, being most frequently eliminated, so that only those would be left which were brown in summer and white in winter. This double selection would be favoured by the fact that there would be, in any case, a change of fur at the beginning of summer; the winter hairs fall out and the fur becomes thinner. The process does not differ essentially from that which takes place in any species when two or more parts or characters, which are not directly connected, have to be changed, such as, for instance, colour and fertility. The struggle for existence will in this case be favourable, on the one hand, to the advantageously coloured, and on the other to the most fertile, and though the two characters may at first only occur separately, they will soon be united by free crossing, until ultimately only those individuals will occur which are at once the most favourably coloured and the most fertile. So in this case there remain only those which are brown in summer and white in winter.

We must ascribe to the influence of the processes of selection the exact regulation of the duration of the winter and summer dress, which has been carefully studied in the case of the variable hare. In the high Alps it remains white for six or seven months, in the south of Norway for eight months, in Northern Norway for nine months, and in Northern Greenland it never loses its white coat at all, as there the snow, even in summer, melts only in some places and for a short time. But apart from concealment there is certainly another adaptation involved here--namely, the growth of the hair as a protection against the cold. From an old experiment made in 1835 by Captain J. Ross, and recently brought to light again by Poulton, we learn that a captive lemming kept in a room in winter did not change colour until it was exposed to the cold. The constitution of animals which become white in winter is thus so organized that the setting in of cold weather acts as a stimulus which incites the skin to the production of white hairs. This predisposition also we must refer to the influence of natural selection, since it must have been very useful to the species that the winter coat should grow just when it was necessary as a protection against cold. This explains at the same time why the predisposition to respond to the stimulus of cold by a growth of winter fur finds expression earlier in those colonies of Arctic animals, such as the hare, which live in Lapland, than in those which live in the south of Norway.

But that it is not the _direct_ influence of cold which colours the hair of a furred animal white we can see from our common hare (_Lepus timidus_), which, in spite of the winter's cold, does not become white, but retains its brown coat, and not less so from the mountain hare (_Lepus variabilis_), which in the south of Sweden also remains brown, although the winter there may be exceedingly cold. But as the covering of the ground with snow is not so uninterrupted there as in the higher North, a white coat would be not a better protection than a brown one, but a worse. The white colouring of Arctic animals is therefore not directly due to the influence of the climate, as has often been maintained, but is due to it indirectly, that is, through the operation of natural selection. I have tried to make this clear by means of this example, so that we may not have to repeat it in considering those which are to follow.

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But all attempts at any other explanation are even more decidedly excluded when we turn our attention to more complicated cases of colour-adaptation, which are not confined to the simple, general coloration, but are helped by markings and colour-patterns, that is, by schemes of colour.

Thus numerous caterpillars exhibit definite lines and spots on their ground-colouring, which, in one way or another, aid in protecting them from their enemies.

The green grass-eating caterpillar of many of our _Satyridæ_ has two or more darker or lighter lines running down the sides of its body, which make it much less conspicuous among the grasses on which it feeds than if it were a uniform green mass (Fig. 2). Not infrequently the colour and form present a remarkably close resemblance to the inflorescences or fruit-ears of the grasses. Caterpillars marked thus are never found on the leaves of trees, where they would immediately catch the eye. It is true that longitudinal striping often occurs on caterpillars which live on other plants besides grass, but as these other plants grow among the grasses the protective efficacy is just the same. This is the case with the Pieridæ (Garden Whites).

All the caterpillars of our Sphingidæ, on the other hand, which live on bushes and trees, have on the sides of the segments light oblique stripes, seven in number, which are disposed to the longitudinal axis of the body at the same angle as the lateral veins of a leaf of their food-plant have to the mid-rib. It cannot of course be said that the caterpillar thereby gains the appearance of a leaf, indeed, if one sees it apart from its food-plant it does not look in the least like a leaf, but among the leaves of a bush or tree this marking secures it in a high degree from discovery. Thus the caterpillar of the eyed hawk-moth (_Smerinthus ocellatus_), when it is sitting among the crowded foliage of a willow, is often very difficult to find, because its large green body does not appear as a single green spot, but is divided by the oblique lateral stripes into sections like the half of a willow leaf, so that even a searching glance is led astray, there being nothing to focus attention on the animal as distinguished from its surroundings (Fig. 3). As a boy I often had the interesting experience of overlooking a caterpillar which was sitting just before me, until after a time I chanced to hit upon the exact spot in the field of vision.

In the majority of these caterpillars with oblique stripes, the likeness to the half of a leaf is heightened by the fact that the light oblique row is accompanied by a broader coloured band, suggesting the shade of the leaf's mid-rib. The caterpillar of _Sphinx ligustri_ has a lilac band, and that of _Sphinx atropos_ a blue one. In both cases it is difficult to believe that such striking colours can secure the animals from discovery, yet among the blending shadows of the leaf-complex of their food-plant they greatly increase their resemblance to a leaf-surface. Of the death's-head caterpillar (_Sphinx atropos_) this sounds almost incredible, for this form is chiefly a bright golden yellow, and the narrow white oblique stripes have sky-blue borders becoming darker towards the under side; but it must not be forgotten that the potato is not the true food-plant of the species, for it lives, in its true home in Africa, and also in the south of Spain, on wild solanaceous plants, which, we are informed by Noll, have precisely these colours--golden-yellow and blue in the blossom, the fruit, and in part also in the leaves and stem. There the caterpillars sit the whole day long on the plants, while with us they have formed the habit of feeding only in the twilight and at night, and concealing themselves in the earth by day, a habit that is found in other caterpillars also, and which we must again ascribe to a process of natural selection.

Some caterpillars exhibit other, more complex markings, which do not protect them by rendering them difficult to detect, but by terrifying the enemy who has discovered them, and warning him away. Such terrifying or aggressive colours are to be found, for instance, in the caterpillars of the Sphingid genus _Chærocampa_ in the form of large eye-like spots, which occur in pairs close together on the fourth and fifth segments of the animal. Children and those unfamiliar with animals take these for true eyes; and as the caterpillar, when it is threatened by an enemy, draws in the head and anterior segments, so that the fourth one is greatly distended, the eye-spots seem to stand on a thick head (Fig. 4), and it cannot be wondered at that the smaller birds, lizards, and other enemies are so terrified that they refrain from attacking. Even hens hesitate to seize such a caterpillar in its defiant attitude, and I once looked on for a long time in a hen-coop while one hen after another rushed to pick up a caterpillar I had placed there, but, when close to it, hastily drew back the head already prepared to strike. Even a gallant cock was a long time in making up his mind to attack the terrible beast, and drew back repeatedly before he at length ventured to strike a deadly blow with his bill. After the first stroke the caterpillar, of course, was lost. Thus even this disguise is only a _relative_ protection, effective only against smaller enemies. But that these are really frightened away, I had once an opportunity of observing, when I put a caterpillar of the common elephant hawk-moth (_Chærocampa elpenor_) in the feeding-trough of a hencoop, and a sparrow flew down to feed from the trough. It descended at first with its back to the caterpillar and fed cheerily. But when by chance it turned round, and spied the caterpillar, it scurried hastily away.

Among Lepidoptera, too, eye-spots often occur on the wings, and to some extent, at least, they have in this case also the significance of warning marks. Take, for instance, the large blue and black eye-spots on the posterior wings of the eyed hawk-moth (_Smerinthus ocellatus_). When the insect is sitting quietly the two spots are not visible, as they are covered by the anterior wings, but as soon as the creature is alarmed it spreads all four wings, and now both eyes stand boldly out on the red posterior wings and alarm the assailant, as they give the impression of the head of a much larger animal (see Fig. 5). There are also eye-like spots which have not this significance and effect, as, for instance, the 'eye-spots' on the train-feathers of the peacock and the Argus pheasant, or the little eye-like spots on the under surface of many diurnal butterflies. In the first case, it is a matter of decoration; in the second, perhaps of the mimicry of dewdrops, which increases still further the resemblance to a withered leaf; but there are undoubtedly many cases in which the eye-spots serve as means of frightening off enemies, and these cases are especially common among butterflies.

Such warning marks are in no way contradictory to the sympathetic colouring of the rest of the body, and indeed we usually find them in combination with it. In some cases the eye-spot, though very conspicuous, is covered, as in the eyed hawk-moth, when at rest, by the sympathetically coloured parts--in this instance the anterior wings. In other cases eye-spots of considerable size lie clearly exposed, but exhibit the same sympathetic colours as the whole of the rest of the wing-surface. In this case they do not interfere with the protective influence of general colouring, because they are only visible from a very short distance. This is the case in the large _Caligo_ species of South America, which only fly for a short time in the early morning and in the evening, remaining concealed throughout the day in dark shadowy places, where the mingled colouring of brown, grey, yellow, and black on the under surfaces of the wings prevents their being recognized from a distance as butterflies at all. But even the best sympathetic colouring is not an absolute protection, and when the insect is discovered by an enemy near at hand, the terrifying mark, a large deep-black spot on the posterior wing, comes into effect, and scares the assailant away.

In such cases the sympathetic colouring was probably the first to arise, and the eye-spot was developed later by a new process of selection, brought about by the necessity of protecting the species more effectively than by mere inconspicuousness alone. In many cases it can be proved that the power of scaring off an enemy did not begin with the formation of the eye-spot, but with the development of a new instinct. When the caterpillar of _Chærocampa elpenor_ is attacked it immediately assumes the defiant attitude described above, but the same striking attitude is assumed by the caterpillars of the allied American genus _Darapsa_, as I learn from an old illustration by Abbot and Smith, although this form possesses no eye-spots (Fig. 7). Thus, then, metaphorically speaking, the caterpillar at first attempted to scare off its enemy by a terrifying attitude alone, and it was only subsequently, in the course of the phyletic evolution, that the eye-spots were added, in the elephant hawk-moths and other species, to heighten the terrifying effect. But that the eye-spot did not make its appearance suddenly is proved by several American species of _Smerinthus_, in which they are much less perfectly developed than in the European species. In these Sphingidæ, too, the defiant attitude was evolved earlier than the eye-spots, as we may see from our poplar hawk-moth (_Smerinthus populi_), which, when alarmed, spreads out all four wings in the same peculiar manner which in the eyed hawk-moth (_Smerinthus ocellatus_) displays the eye-spots; it strikes about with its wings as if to scare off the enemy, an effect which will certainly be more surely achieved if, at the same time, a pair of eyes becomes suddenly visible.

Sympathetically coloured caterpillars are, however, by no means the only ones; there are some with such striking, glaring colours that, far from rendering their possessors inconspicuous, they make them visible from a long way off; but this apparent contradiction of the theory of the colour-adaptation of animals that require protection has been explained by the acuteness of Alfred Russel Wallace. We know that among insects, and also among caterpillars, there are many which have a repulsive taste. In any case, certain caterpillars are rejected by many birds and lizards. Such species are, therefore, relatively safe from being devoured. If they were protectively coloured, or if, moreover, they resembled caterpillars with an agreeable taste, they would gain little advantage from their unpalatability; for the birds would at first take them for eatable, and would only discover their repulsiveness on attempting to eat them. But a caterpillar which has received a single stroke from a bird's bill is doomed to death. It must therefore be of the greatest advantage for unpalatable caterpillars, and unpalatable animals generally, to be in their colouring as conspicuously distinguishable as possible from the edible species. Hence, then, the glaring colours, which we can now refer without any further difficulty to the process of natural selection, for every individual of an ill-tasting species that is more conspicuously coloured than its fellows must have an advantage over them, and must have a better chance of surviving, because it will be less easily mistaken for a member of an edible species.

I should like to discuss one other phenomenon, which is well calculated to give us a deeper insight into the transformation processes of organisms--I refer to the remarkable dimorphism of colour which occurs in many of the species of caterpillar just described.

The caterpillar of the convolvulus hawk-moth (_Sphinx convolvuli_) is in its full-grown stage green, like the wild convolvulus on which it lives, or brown like the ground on which its food-plant grows. It thus shows a double adaptation, each of which is capable of protecting it to a certain extent, and we might think to the _same_ extent. But that is not so, the brown colouring is a more effective protection than the green, as we may learn from two facts. In the first place, the four young stages of the caterpillar are green, and it only becomes brown in the last stage, though sometimes even then it remains green. This shows that the brown is a relatively modern adaptation, and it could not have arisen had it not been better than the original green. In the second place, the green-coloured caterpillars of the convolvulus hawk-moth are nowadays much less numerous than the brown ones, and this implies that the latter survive oftener in the struggle for existence. We have here an interesting case of an easily recognizable process of selection still going on between the old green and the newer brown variety.