Chapter X: Part 10

But, while only of late this region has had a public history, the long-forgotten years between the Glacial period and the expedition of Lewis and Clarke were not without interest in the history of the trout. For all these years the fishes have been trying to mount the waterfalls in order to ascend to the plateau above. Year after year, as the spawning-time came on, they leaped against the falls of the Gardiner, the Gibbon, and the Firehole Rivers, but only to fall back impotent in the pools at their bases. But the mightiest cataract of all, the great falls of the Yellowstone, they finally conquered, and in this way it was done: not by the trout of the Yellowstone River, but by their brothers on the other side of the Divide. These followed up the Columbia to the head-waters of the Snake River, its great tributary, past the beautiful Heart Lake, and then on to the stream now called Pacific Creek, which rises on the very crest of the Divide. In the space between this stream, which flows west to help form the Snake River, and a smaller stream now called Atlantic Creek, flowing down the east slope of the Divide, the great chain of the Rocky Mountains shrinks to a narrow plateau of damp meadow, not a fourth of a mile in width; and some years, when the snows are heavy and melt late in the spring, this whole region is covered with standing water. The trout had bided their time until they found it so, and now they were ready for action. Before the water was drained they had crossed the Divide and were descending on the Atlantic side toward the Yellowstone Lake. As the days went by, this colony of bold trout spirits grew and multiplied and filled the waters of the great clear lake, where their descendants remain to this day. And no other fishes--not the chub, nor the sucker, nor the white-fish, nor the minnow, nor the blob--had ever climbed Pacific Creek. None of them were able to follow where the trout had gone, and none of them have ever been seen in the Yellowstone Lake. What the trout had done in this lake--their victories and defeats, their struggles with the bears and pelicans, and with the terrible worm, joint enemy of trout and pelicans alike--must be left for another story.

So the trout climbed the Yellowstone Falls by way of the back staircase. For all we know, they have gone down it on the other side. And in a similar way, by stealing over from Blacktail Deer Creek, they overcame the Undine Falls in Lava Creek and passed its steep obsidian walls, which not all the fishes in the world could climb.

In the Gibbon River the cataracts have proved to the trout an impassible barrier; but, strangely enough, its despised associate, the sluggish, chunky blob, the little soft-bodied, smooth, black tadpole-like fellow, with twinkling eyes and a voracious appetite--a fish who cannot leap at all--has crossed this barrier. Hundreds of blob live under the stones in the upper reaches of the stream, the only fish in the Gibbon waters. There he is, and it is a standing puzzle even to himself to know how he got there. We might imagine, perhaps, that some far-off ancestor, some ancient Queen of the Blobs, was seized by an osprey and carried away in the air. Perhaps an eagle was watching and forced the osprey to give up its prey. Perhaps in the struggle the blob escaped, falling into the river above the falls, to form the beginning of the future colony. At any rate, there is the great impassable waterfall, the blob above it and below. The osprey has its nest on a broken pine tree, above the cataract, and its tyrant master, the bald eagle, watches it from a still higher crag whenever it goes fishing.

Two years ago the Hon. Marshall McDonald, whose duty as United States Fish Commissioner it is to look after the fishes wherever they may be, sent me to this country to see what could be done for his wards. It was a proud day when I set out from Mammoth Hot Springs astride a black cayuse, or Indian pony, which answered to the name of Jump, followed by a long train of sixteen other cayuses of every variety of color and character, the most notable of all being a white pony called Tinker. At some remote and unidentified period of her life she had bucked and killed a tradesman who bestrode her against her will, and thereby, as in the old Norse legends, she has inherited his strength, his wickedness, and his name. And when, after many adventures, I came back from this strange land and told the story of its fishes other men were sent out from Washington with nets and buckets. They gathered up the trout and carried them to the rivers above the falls, and now all the brooks and pools of the old lava-bed, the fairest streams in the world, are full of their natural inhabitants.

THE COLORS OF ANIMALS

(FROM CHAPTERS IN POPULAR NATURAL HISTORY.)

BY SIR JOHN LUBBOCK, BART., M.P., F.B.S., ETC.

The color of animals is by no means a matter of chance; it depends on many considerations, but in the majority of cases tends to protect the animal from danger by rendering it less conspicuous. Perhaps it may be said that if coloring is mainly protective, there ought to be but few brightly colored animals. There are, however, not a few cases in which vivid colors are themselves protective. The kingfisher itself, though so brightly colored, is by no means easy to see. The blue harmonizes with the water, and the bird as it darts along the stream looks almost like a flash of sunlight; besides which, protection is not the only consideration. Let us now consider the prevalent colors of animals and see how far they support the rule.

Desert animals are generally the color of the desert. Thus, for instance, the lion, the antelope, and the wild ass are all sand-colored. "Indeed," says Canon Tristram, "in the desert, where neither trees, brushwood, nor even undulation of the surface afford the slightest protection to its foes, a modification of color 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 sand color."

It is interesting to note that, while the lion is sand-colored like the desert, the long, upright, yellow stripes of the tiger make it very difficult to see the animal among the long dry grasses of the Indian jungles in which it lives. The leopard, again, and other tree cats are generally marked with spots which resemble gleams of light glancing through the leaves.

The colors of birds are in many cases perhaps connected with the position and mode of construction of their nests. Thus, we know that hen birds are generally less brightly colored than the cocks, and this is partly, perhaps, because bright colors would be a danger to the hens while sitting on their eggs. When the nest is placed underground or in the hole of a tree, etc., we find it no longer to be such an invariable rule that the hen bird is dull-colored; but, on the contrary, she is then often as gaily colored as the male. Such, for instance, is the case with the hen kingfisher, which is one of the brightest of British birds and one of the very few which make their nests underground; the hen woodpecker, which is also gaily colored and builds in hollow trees, forms a second instance.

In the few cases where the hens are as conspicuously colored as the cocks, and yet the nest is open to view, we generally find that the hens are strong, pugnacious birds, and well able to defend themselves. There are even instances, though these are comparatively rare, in which the hens are more brilliantly colored than the cocks; and it is an interesting fact that it is then the cocks, and not the hens, which hatch the eggs.

It therefore seems to be a rule, with very few exceptions, that when both the cocks and hens are of strikingly gay or conspicuous colors, the nest is such as to conceal the sitting bird; while, whenever there is a striking contrast of colors, the nest is open and the sitting bird exposed to view.

Again, most fishes are dark above and pale below. This points to the same fact, for when one looks down into the dark water, the dark color of their backs renders them the less easy to distinguish; while, to an enemy looking up from below, the pale belly would be less conspicuous against the light of the sky. Those fishes which live deep down in the depths of the ocean present no such contrast between the upper and under surface. Many of the smaller animals which live in the sea are as transparent as glass, and are consequently very difficult to distinguish.

It is sometimes said that if animals were really colored with reference to concealment, sheep would be green, like grass. This, however, is quite a mistake. If they were green they would really be more easy to see. In the gray of the morning and the evening twilight, just the time when wild animals generally feed, gray and stone colors are most difficult to distinguish. Sheep were originally mountain animals, and every one who has ever been on a mountain-side knows how difficult it is to distinguish a sheep, at some distance, from a mass of stone or rock.

It is, again, a great advantage to the rabbit and hare to be colored like earth; black or white rabbits are more easy to see, and consequently more likely to be killed. This, however, does not apply to those which are kept in captivity, and we know that tame rabbits are often black and white. Again, in the far north, where for months together the ground is covered with snow, the white color, which would be a danger here, becomes an advantage; and many Arctic animals, like the polar bear and polar hare, are white, while others, such as the mountain hare and ptarmigan, change their color, being brown in summer and white in winter. So are the Arctic fox and the ermine, to whom it is then an advantage to be white, not to avoid danger, but in order that they may be the more easily able to steal unperceived upon their prey.

Many of the cases in which certain insects escape danger by their similarity to plants are well known; the leaf insect and the walking-stick insect are familiar and most remarkable cases. The larvae of insects afford, also, many interesting examples, and in other respects teach us, indeed, many instructive lessons. It would be a great mistake to regard them as merely preparatory stages in the development of the perfect insect. They are much more than this, for external circumstances act on the larvae, as well as on the perfect insect: both, therefore, are liable to adaptation. In fact, the modifications which insect larvae undergo may be divided into two kinds--developmental, or those which tend to approximation to the mature form; and adaptational or adaptive, those which tend to suit them to their own mode of life.

It is a remarkable fact, that the forms of larvae do not depend on those of the mature insect. In many cases, for instance, very similar larvae produce extremely dissimilar insects. In other cases, similar, or comparatively similar, perfect insects have very dissimilar larvae. Indeed, a classification of insects founded on larva would be quite different from that founded on the perfect insects. The group to which the bees, wasps, and ants belong, for instance, and which, so far as the perfect insects are concerned, form a very natural division, would be divided into two; or rather one portion of them--namely, the saw-flies--would be united to the butterflies and moths. Now, why do the larvae of saw-flies differ from those of their allies, and resemble those of butterflies and moths? It is because their habits differ from those of ants and bees, and they feed on leaves like ordinary caterpillars.

In some cases the form changes considerably during the larval state. From this point of view, the transformations of a small beetle, called _Sitaris_, which have been carefully observed by M. Fabre, are peculiarly interesting.

The genus Sitaris, which is allied to the blister-fly and to the oil-beetle, is parasitic on a kind of solitary bee which excavates subterranean galleries, each leading to a cell. The eggs of the beetle, which are deposited at the entrance of the galleries made by the bees, are hatched at the end of September or beginning of October, and we might not unnaturally expect that the young larvae, which are active little creatures with six serviceable legs, would at once eat their way into the cells of the bee. No such thing: till the month of April following they remain without leaving their birthplace, and consequently without food; nor do they in this long time change either in form or size. M. Fabre ascertained this, not only by examining the burrow of the bees, but also by direct observations of some young larvae kept in captivity. In April, however, his captives at last awoke from their long lethargy, and hurried anxiously about their prisons. Naturally inferring that they were in search of food, M. Fabre supposed that this would consist either of the larvae or pupae of the bee, or of the honey with which it stores its cell. All three were tried without success. The first two were neglected; and the larvae, when placed on the latter, either hurried away or perished in the attempt, being evidently unable to deal with the sticky substance. M. Fabre was in despair. The first ray of light came to him from our countryman Newport, who ascertained that a small parasite found on one of the wild bees was, in fact, the larva of the oil-beetle. The larvae of Sitaris much resembled this larva. Acting on this hint, M. Fabre examined many specimens of the bee, and found on them at last the larvae of his Sitaris. The males of the bee emerge from the pupae sooner than the females, and M. Fabre ascertained that, as they come out of their galleries, the little Sitaris larvae fasten upon them. Not, however, for long: instinct teaches them that they are not yet in the straight path of development; and, watching their opportunity, they pass from the male to the female bee. Guided by these indications, M. Fabre examined several cells of the bee; in some, the egg of the bee floated by itself on the surface of the honey; in others, on the egg, as on a raft, sat the still more minute larva of the Sitaris. The mystery was solved. At the moment when the egg is laid, the Sitaris larva springs upon it. Even while the poor mother is carefully fastening up her cell, her mortal enemy is beginning to devour her offspring; for the egg of the bee serves not only as a raft, but as a repast. The honey, which is enough for either, would be too little for both; and the Sitaris, therefore, at its first meal, relieves itself from its only rival. After eight days the egg is consumed, and on the empty shell the Sitaris undergoes its first transformation, and makes its appearance in a very different form.

The honey, which was fatal before, is now necessary--the activity, which before was necessary, is now useless; consequently, with the change of skin, the active, slim larva changes into a white fleshy grub, so organized as to float on the surface of the honey, with the mouth beneath and the breathing-holes above the surface; for insects breathe, not as we do through the mouth, but through a row of holes arranged along the side. In this state it remains until the honey is consumed; then the animal contracts, and detaches itself from its skin, within which the further transformations take place. In the next stage the larva has a solid corneous envelope and an oval shape, and, in its color, consistency, and immobility, resembles the chrysalis of a fly. The time passed in this condition varies much. When it has elapsed, the animal moults again, again changes its form; after this, it becomes a pupa, without any remarkable peculiarities. Finally, after these wonderful changes and adventures, in the month of August the perfect beetle makes its appearance.

In fact, whenever in any group we find differences in form or color, we shall always find them associated with differences in habit. Let us take the case of Caterpillars. The prevailing color of caterpillars is green, like that of leaves. The value of this to the young insect, the protection it affords, are obvious. We must all have observed how difficult it is to distinguish small green caterpillars from the leaves on which they feed. When, however, they become somewhat larger, their form betrays them, and it is important that there should be certain marks to divert the eye from the outlines of the body. This is effected, and much protection is given, by longitudinal lines (Fig. 1), which accordingly are found on a great many caterpillars. These lines, both in color and thickness, much resemble some of the lines on leaves (those, for instance, of grasses), and also the streaks of shadow which occur among foliage. If this be the explanation of them, then they ought to be wanting, as a general rule, in very small caterpillars, and should prevail most among those which feed on or among grasses.

Now, similar lines occur on a great number of caterpillars belonging to most different groups of butterflies and moths, as you may see by turning over the illustrations of any monograph of the group. They exist among the Hawk-moths--as, for instance, in the Humming-bird Hawk-moth; they occur in many butterflies, especially in those which feed on grass; and in many moths. But you will find that the smallest caterpillars rarely possess these white streaks. As regards the second point, also, the streaks are generally wanting in caterpillars which feed on large-leaved plants. The _Satyridae_, on the contrary, all possess them, and all live on grass. In fact we may say, as a general rule, that these longitudinal streaks only occur on caterpillars which live on or among narrow-leaved plants. As the insect grows, these lines often disappear on certain segments, and are replaced by diagonal lines. These diagonal lines (Fig. 2) occur in a great many caterpillars, belonging to the most distinct families of butterflies and moths. They come off just at the same angle as the ribs of leaves, and resemble them very much in general effect. They occur also especially on species which feed on large-leaved plants; and I believe I may say that though a great many species of caterpillars present these lines, they rarely, if ever, occur in species which live on grass; while, on the contrary, they are very frequent in those species which live on large-leaved plants.

It might at first be objected to this view that there are many cases, as in the Elephant-Hawk-moth, in which caterpillars have both. A little consideration, however, will explain this. In small caterpillars these oblique lines would be useless, because they must have some relation, not only in color, but in their distance apart, to the ribs of the leaves. Hence, while there are a great many species which have, longitudinal lines when young, and diagonal ones when they are older and larger, there is not, I believe, a single one which begins with diagonal lines, and then replaces them with longitudinal ones. The disappearance of the longitudinal lines on those segments which have diagonal ones, is striking, where the lines are marked. It is an advantage, because white lines crossing one another at such an angle have no relation to anything which occurs in plants, and would make the creature more conspicuous. When, therefore, the diagonal lines are developed, the longitudinal ones often disappear. There is one other point in connection with these diagonal lines to which I must call your attention.

In many species they are white, but in some cases--as, for instance, in the beautiful green caterpillar of the Privet-Hawk-moth--the white streak is accompanied by a colored one, in that case lilac. At first we might think that this would be a disadvantage, as tending to make the caterpillar more conspicuous; and in fact, if we put one in full view--for instance, out on a table--and focus the eye on it, the colored lines are very striking. But we must remember that the habit of the insect is to sit on the lower side of the leaf, generally near the middle rib, and in the subdued light of such a situation, especially if the eye be not looking exactly at them, the colored lines beautifully simulate a line of soft shadow, such as must always accompany a strong rib; and I need not tell any artist that the shadows of yellowish-green must be purplish. Moreover, any one who has ever found one of these large caterpillars will, I am sure, agree with me that it is surprising, when we consider their size and conspicuous coloring, how difficult it is to see them.

But though the prevailing color of caterpillars is green, there are numerous exceptions. In one great family of moths the prevailing color is brown. These caterpillars, however, escape observation by their great similarity to brown twigs--a resemblance which is heightened by their peculiar attitudes, and in many cases by the existence of warts or protuberances, which look like buds. Some, however, even of these caterpillars, when very young, are green. Again, some caterpillars are white. These feed on and burrow in wood. The Ringlet Butterfly also has whitish caterpillars, and this may at first sight appear to contradict the rule, since it feeds on grass. Its habit is, however, to keep at the roots by day, and feed only at night.

In various genera we find Black caterpillars, which are of course very conspicuous, and, so far as I know, not distasteful to birds. In such cases, however, it will be found that they are covered with hairs or spines, which protect them from most birds. In these species the bold dark color may be an advantage, by rendering the hair more conspicuous. Many caterpillars are black and hairy, but I do not know any large caterpillar which is black and smooth.

Brown caterpillars, also, are frequently protected by hairs or spines in the same way; but, unlike black ones, they are frequently naked. These fall into two principal categories: firstly, those which, like the Geometridae, put themselves into peculiar and stiff attitudes, so that in form, color, and position they closely resemble bits of dry stick; and, secondly, those which feed on low plants, concealing themselves on the ground by day, and only coming out in the dark.

Yellow and yellowish-green caterpillars are abundant, and their color is a protection. Red and blue, on the contrary, are much less common colors, and are generally present as spots.

Moreover, caterpillars with red lines or spots are generally hairy, and this for the reason given above. Such species, therefore, would be avoided by birds. There are, no doubt, some apparent exceptions. The Swallow-tail Butterfly, for instance, has red spots and still is smooth; but as it emits a strongly-scented liquid when alarmed, it is probably distasteful to birds. I cannot recall any other case of a British caterpillar which has conspicuous red spots or lines, and yet is smooth.

Blue is, among caterpillars, even a rarer color than red. Indeed, among our larger larvae, the only cases I can recall are the Lappets, which have two conspicuous blue bands, the Death's-head Moth, which has broad diagonal bands, and two of the Hawk-moths, which have two bright blue oval patches on the third segment. The Lappets are protected by being hairy, but why they have the blue bands I have no idea. It is interesting, that both the other species frequent plants which have blue flowers. The peculiar hues of the Death's-head caterpillar, which feeds on the potato, unite so beautifully the brown of the earth, the yellow and green of the leaves, and the blue of the flowers, that, in spite of its size, it can scarcely be perceived unless the eye be focussed exactly upon it.

The Oleander Hawk-moth is also an interesting case. Many of the Hawk-moth caterpillars have eye-like spots, to which I shall have to allude again presently. These are generally reddish or yellowish, but in this species, which feeds on the periwinkle, they are bright blue, and in form as well as color closely resemble the blue petals of that flower. One other species, the Sharp-winged Hawk-moth, also has two smaller blue spots, with reference to which I can make no suggestion. It is a very rare species, and I have never seen it. Possibly, in this case, the blue spots may be an inherited character, and have no reference to the present habits. They are, at any rate, quite small.

No one who looks at any representations of Hawk-moth caterpillars can fail to be struck by the peculiar coloring of those belonging to the Pine Moth, which differ in style of coloring from all other sphinx larvae, having longitudinal bands of brown and green. Why is this? Their _habitat_ is different. They feed on the leaves of the pinaster, and their peculiar coloring offers a general similarity to the brown twigs and narrow green leaves of a conifer. There are not many species of butterflies or moths which feed on the pine, but there are a few: and most, if not all of them, have a very analogous style of coloring to that of the Pine Moth, while the latter has also tufts of bluish-green hair which singularly mimic the leaves of the pine. It is still more remarkable that in a different order of insects we again find species--for instance one of the saw-flies--which live on the pine, and in which the same style of coloring is repeated.

Let us now take a single group, and see how far we can explain its various colors and markings, and what are the lessons which they teach us. For this purpose, I think I cannot do better than select the larvae of the Hawk-moths, which have just been the subject of a masterly work by Dr. Weissmann, from which most of the following facts are taken.

The caterpillars of this group are very different in color--green, white, yellow, brown, sometimes even gaudy, varied with spots, patches, streaks, and lines. Now, are these differences merely casual and accidental, or have they a meaning and a purpose? In many, perhaps in most cases, the markings serve for the purpose of concealment. When, indeed, we see caterpillars represented on a white sheet of paper, or if we put them on a plain table, and focus the eye on them, the colors and markings would seem, if possible, to render them even more conspicuous; but amongst the intricate lines and varied colors of foliage and flowers, and if the insect be a little out of focus, the effect is very different.

Let us begin with the Elephant Hawk-moth. The caterpillars (Fig. 3), as represented in most entomological works, are of two varieties, most of them brown, but some green. Both have a white line on the three first segments; two remarkable eye-like spots on the fourth and fifth, and a very faint median line; and are rather more than four inches long. I will direct your attention specially, for the moment, to three points:--What do the eye-spots and the faint lateral line mean? and why are some green and some brown, offering thus such a marked contrast to the leaves of the small epilobe on which they feed? Other questions will suggest themselves later. I must now call your attention to the fact, that when the caterpillars first quit the egg, and come into the world (Fig. 4), they are quite different in appearance, being, like so many other small caterpillars, bright green, and almost exactly the color of the leaves on which they feed. That this color is not the necessary or direct consequence of the food, we see from the case of quadrupeds, which, as I need scarcely say, are never green. It is, however, so obviously a protection to small caterpillars, that this explanation of their green color suggests itself to every one.

After five or six days, and when they are about a quarter of an inch in length, they go through their first moult. In their second stage (Fig. 5), they have two white lines, stretching along the body from the horn to the head; and after a few days (Fig. 6), but not at first, traces of the eye-spots appear on the fourth and fifth segments, shown by a slight wave in the upper line. After another five or six days, and when about half an inch in length, our caterpillars moult again. In their third stage (Fig. 7), the commencement of the eye-spots is more marked, while, on the contrary, the lower longitudinal line has disappeared. After another moult (Fig. 8), the eye-spots are still more distinct, the white gradually becomes surrounded by a black line, while in the next stage (Fig. 9) the centre becomes somewhat violet. The white lines have almost or entirely disappeared, and in some specimens faint diagonal lines make their appearance. Some few assume a brownish tint, but not many. A fourth moult takes place in seven or eight days, and when the caterpillars are about an inch and a half in length. Now, the difference shows itself still more between the two varieties, some remaining green, while the majority become brown. The eye-spots are more marked, and the pupil more distinct, the diagonal lines plainer, while the white line is only indicated on the first three, and on the eleventh segment. The last stage (Fig. 9) has been already described.

Now, the principal points to which I wish to draw attention are (1) the green color, (2) the longitudinal lines, (3) the diagonal lines, (4) the brown color, and (5) the eye-spots.

As regards the first three, however, I think I need say no more. The value of the green color to the young larva is obvious; nor is it much less clear that when the insect is somewhat larger, the longitudinal lines are a great advantage, while subsequently diagonal ones become even more important.

The next point is the color of the mature caterpillars. We have seen that some are green, and others brown. The green ones are obviously merely those which have retained their original color. Now for the brown color. This probably makes the caterpillar even more conspicuous among the green leaves than would otherwise be the case. Let us see, then, whether the habits of the insect will throw any light upon the riddle. What would you do if you were a big caterpillar? Why, like most other defenceless creatures, you would feed by night, and lie concealed by day. So do these caterpillars. When the morning light comes, they creep down the stem of the food plant, and lie concealed among the thick herbage, and dry sticks and leaves, near the ground; and it is obvious that under such circumstances the brown color really becomes a protection. It might indeed be argued that the caterpillars, having become brown, concealed themselves on the ground; and that we were, in fact, reversing the state of things. But this is not so; because, while we may say, as a general rule, that (with some exceptions due to obvious causes) large caterpillars feed by night and lie concealed by day, it is by no means always the case that they are brown; some of them still retaining the green color. We may then conclude that the habit of concealing themselves by day came first, and that the brown color is a later adaptation. It is, moreover, interesting to note that while the caterpillars which live on low plants often go down to the ground and turn brown, those which feed on large trees or plants remain on the under side of the leaves, and retain their green color.

Thus, in the Eyed Hawk-moth, which feeds on the willow and sallow; the Poplar Hawk-moth, which feeds on the poplar; and the Lime Hawk-moth, which frequents the lime, the caterpillars all remain green; while in those which frequent low plants, such as the Convolvulus Hawk-moth, which frequents the convolvulus; the Oleander Hawk-moth, which feeds in this country on the periwinkle; and other species, most of the caterpillars turn brown. There are, indeed, some caterpillars which are brown, and still do not go down to the ground--as, for instance, those of the Geometridae generally. These caterpillars, however, as already mentioned, place themselves in peculiar attitudes, which, combined with their brown color, make them look almost exactly like bits of stick or dead twigs.

The last of the five points to which I called your attention was the eye-spots. In some cases, spots may serve for concealment, by resembling the marks on dead leaves. In one species, which feeds on the hippophae, or sea buckthorn, a gray-green plant, the caterpillar also is a similar gray-green, and has, when full grown, a single red spot on each side--which, as Weissmann suggests, at first sight much resembles in color and size one of the berries of the hippophae. This might, at first, be supposed to constitute a danger, and therefore to be a disadvantage; but the seeds, though present, are not ripe, and consequently are not touched by birds. Again, in another caterpillar, there is an eye-spot on each segment, which mimics the flower of the plant on which it feeds. White spots, in some cases, also resemble the spots of light which penetrate foliage. In other instances, however, and at any rate in our Elephant Hawk-moth, the eye-spots certainly render the insect more conspicuous.

Now in some cases, this is an advantage, rather than a drawback. Suppose that from the nature of its food, from its being covered with hair, or from any other cause, a small green caterpillar were very bitter, or disagreeable or dangerous as food, still, in the number of small green caterpillars which birds love, it would be continually swallowed by mistake. If, on the other hand, it had a conspicuous and peculiar color, its evil taste would serve to protect it, because the birds would soon recognize and avoid it, as has been proved experimentally. I have already alluded to a case of this among the Hawk-moths, in a species which, feeding on euphorbia, with its bitter milky juice, is very distasteful to birds, and is thus actually protected by its bold and striking colors. The spots on our Elephant Hawk-moth caterpillar do not admit of this explanation, because the insect is quite good to eat--I mean, for birds. We must, therefore, if possible, account for these spots in some other way. There can, I think, be little doubt that Weissmann is right when he suggests that the eye-spots actually protect the caterpillar, by frightening its foes.

Every one must have observed that these large caterpillars--as, for instance, that of the small Elephant Hawk-moth (Fig. 10)--have a sort of uncanny poisonous appearance; that they suggest a small thick snake or other evil beast, and the so-called "eyes" do much to increase the deception. Moreover, the segment on which they are placed is swollen, and the insect, when in danger, has the habit of retracting its head and front segments, which gives it an additional resemblance to some small reptile. That small birds are, as a matter of fact, afraid of these caterpillars (which, however, I need not say, are in reality altogether harmless), Weissmann has proved by actual experiment. He put one of these caterpillars in a tray in which he was accustomed to place seed for birds. Soon a little flock of sparrows and other small birds assembled to feed as usual. One of them lit on the edge of this tray, and was just going to hop in, when she spied the caterpillar. Immediately she began bobbing her head up and down, but was afraid to go nearer. Another joined her, and then another, until at last there was a little company of ten or twelve birds, all looking on in astonishment, but not one ventured into the tray; while one bird, which lit in it unsuspectingly, beat a hasty retreat in evident alarm, as soon as she perceived the caterpillar. After watching for some time, Weissmann removed it, when the birds soon attacked the seeds. Other caterpillars also are probably protected by their curious resemblance to spotted snakes.

Moreover, we may learn another very interesting lesson from these caterpillars. They leave the egg, as we have seen, a plain green, like so many other caterpillars, and gradually acquire a succession of markings, the utility of which I have just attempted to explain. The young larva, in fact, represents an old form, and the species, in the lapse of ages, has gone through the stage which each individual now passes through in a few weeks. Thus, the caterpillar of _Chaerocampa porcellus_, a species very nearly allied to the Elephant Hawk-moth, passes through almost exactly the same stages as that species. But it leaves the egg with a subdorsal line, which the caterpillar of the Elephant Hawk-moth does not acquire until after its first moult. No one can doubt, however, that there was a time when the new-born caterpillars of the small Elephant Hawk-moth were plain green, like those of the large one. Again, if we compare the mature caterpillars of this group of Hawk-moths, we shall find there are some forms which never develop eye-spots, but which, even when full grown, correspond to the second stage of the Elephant Hawk-moth. Here, then, we seem to have species still in the stage which the Elephant Hawk-moth must have passed through long ago.

The genus _Deilephila_, of which we have three species--the Euphorbia Hawk-moth, the Galium Hawk-moth, and the Rayed Hawk-moth--is also very instructive. The caterpillar of the Euphorbia Hawk-moth begins life of a clear green color, without a trace of the subsequent markings. After the first moult, however, it has a number of black patches, a white line, and a series of white dots, and has, therefore, at one bound, acquired characters which in the Elephant Hawk-moth, as we have seen, were only very gradually assumed. In the third stage, the line has disappeared, leaving the white spots. In the fourth, the caterpillars have become very variable, but are generally much darker than before, and have a number of white dots under the spots. In the fifth stage, there is a second row of white spots under the first. The caterpillars not being good to eat, there is, as has been already pointed out, no need for, or attempt at, concealment. Now if we compare the mature caterpillars of other species of the genus, we shall find that they represent phases in the development of the Euphorbia Hawk-moth. The Sea Buckthorn Hawk-moth, for instance, even when full grown, is a plain green, with only a trace of the line, and corresponds, therefore, with a very early stage of the Euphorbia Hawk-moth; there is another species found in South Russia, which has the line, and represents the second stage of the Euphorbia Hawk-moth; another has the line and the row of spots, and represents, therefore, the third stage; lastly, there are some which have progressed further, and lost the longitudinal line, but they never acquire the second row of spots which characterizes the last stage of the Euphorbia Hawk-moth.

Thus, then, the individual life of certain caterpillars gives us a clue to the history of the species in past ages.

For such inquiries as this, the larvae of Lepidoptera are particularly suitable, because they live an exposed life; because the different species, even of the same genus, often feed on different plants, and are therefore exposed to different conditions; and last, not least, because we know more about the larvae of the butterflies and moths than about those of any other insects. The larvae of ants all live in the dark; they are fed by the perfect ants, and being therefore all subject to very similar conditions, are all very much alike. It would puzzle even a good naturalist to determine the species of an ant larva, while, as we all know, the caterpillars of butterflies and moths are as easy to distinguish as the perfect insects; they differ from one another as much as, sometimes more than, the butterflies and moths themselves.

There are five principal types of coloring among caterpillars. Those which live inside wood, or leaves, or underground, are generally of a uniform pale hue; the small leaf-eating caterpillars are green, like the leaves on which they feed. The other three types may, to compare small things with great, be likened to the three types of coloring among cats. There are the ground cats, such as the lion or puma, which are brownish or sand color, like the open places they frequent. So also caterpillars which conceal themselves by day at the roots of their food-plant, tend, as we have seen, even if originally green, to assume the color of earth. Nor must I omit to mention the _Geometridae_, to which I have already referred, and which, from their brown color, their peculiar attitudes, and the frequent presence of warts or protuberances, closely mimic bits of dry stick. That the caterpillars of these species were originally green, we may infer from the fact that some of them at least are still of that color when first born.

Then there are the spotted or eyed cats, such as the leopard, which live among trees; and their peculiar coloring renders them less conspicuous by simulating spots of light which penetrate through foliage. So also many caterpillars are marked with spots, eyes, or patches of color. Lastly, there are the jungle cats, of which the tiger is the typical species, and which have stripes, rendering them very difficult to see among the brown grass which they frequent. It may, perhaps, be said that this comparison fails, because the stripes of tigers are perpendicular, while those of caterpillars are either longitudinal or oblique. This, however, so far from constituting a real difference, confirms the explanation; because in each case the direction of the lines follows that of the foliage. The tiger, walking horizontally on the ground, has transverse bars; the caterpillar, clinging to the grass in a vertical position, has longitudinal lines; while those which live on large-veined leaves have oblique lines, like the oblique ribs of the leaves.

Red and blue are rare colors among caterpillars. Omitting minute dots, we have six species more or less marked with red or orange. Of these, two are spiny, two hairy, and one protected by scent-emitting tentacles. The orange medio-dorsal line of the Bedford Butterfly is not very conspicuous, and has been omitted in some descriptions. Blue is even rarer than red; in fact, none of our butterfly larvae can be said to exhibit this color.

Now let us turn to the moths. I have taken all the larger species, amounting to rather more than one hundred and twenty; out of which sixty-eight are hairy or downy; and of these forty-eight are marked with black or gray, fifteen brown or brownish, two yellowish-green, one bluish-gray, one striped with yellow and black, and one reddish-gray. There are two yellowish-green hairy species, which might be regarded as exceptions: one, that of the Five-spotted Burnet-moth, is marked with black and yellow, and the other is variable in color, some specimens of this caterpillar being orange. This last species is also marked with black, so that neither of these species can be considered of the green color which serves as a protection. Thus, among the larger caterpillars, there is not a single hairy species of the usual green color. On the other hand, there are fifty species with black or blackish caterpillars, and of these forty-eight are hairy or downy.

In ten of our larger moths the caterpillars are more or less marked with red. Of these, three are hairy, one is an internal feeder, four have reddish lines, which probably serve for protection by simulating lines of shadow, and one, the Euphorbia Hawk-moth, is inedible. The last, the striped Hawk-moth, is rare, and I have never seen the caterpillar; but to judge from figures, the reddish line and spots would render it, not more, but less conspicuous amongst the low herbage which it frequents.

Seven species only of the larger moths have any blue; of these, four are hairy, the other three are Hawk-moths. In one, the Death's Head, the violet color of the side stripes certainly renders the insect less conspicuous among the flowers of the potato, on which it feeds. In the Oleander Hawk-moth there are two blue patches, which, both in color and form, curiously resemble the petals of the periwinkle, on which it feeds. In the third species, the small Elephant Hawk-moth, the bluish spots form the centres of the above-mentioned eye-like spots.

In one family, as already mentioned, the caterpillars are very often brown, and closely resemble bits of stick, the similarity being much increased by the peculiar attitudes they assume. On the other hand, the large brown caterpillars of certain Hawk-moths are night feeders, concealing themselves on the ground by day; and it is remarkable that while those species, such as the Convolvulus Hawk-moth, which feed on low plants, turn brown as they increase in age and size, others, which frequent trees, and cannot therefore descend to the ground for concealment, remain green throughout life. Omitting these, there are among the larger species, seventeen which are brown, of which twelve are hairy, and two have extensile caudal filaments. The others closely resemble bits of stick, and place themselves in peculiar and stiff attitudes.

And thus, summing up the caterpillars, both of butterflies and moths, out of eighty-eight spiny and hairy species, only one is green, and even this may not be protectively colored, since it has conspicuous yellow warts. On the other hand, a very great majority of the black and brown caterpillars, as well as those more or less marked with blue and red, are either hairy or spiny, or have some special protection.

Here, then, I think we see reasons, for many at any rate, of the variations of color and markings in caterpillars, which at first sight seem so fantastic and inexplicable. I should, however, produce an impression very different from that which I wish to convey, were I to lead you to suppose that all these varieties have been explained, or are understood. Far from it; they still offer a large field for study; nevertheless, I venture to think the evidence now brought forward, however imperfectly, is at least sufficient to justify the conclusion that there is not a hair or a line, not a spot or a color, for which there is not a reason--which has not a purpose or a meaning in the economy of nature.

PROTECTIVE RESEMBLANCES IN SPIDERS

BY ELIZABETH G. PECKHAM.[10]

[10] Abbreviated from the occasional Papers of the Natural History Society of Wisconsin, Vol. I., 1889. By permission.

There are, among spiders, two forms of protective modification: the first, including all cases of protective resemblance to vegetable and inorganic things--that is, all modifications of color or of color and form that tend to make them inconspicuous in their natural relations--I shall call direct protection. The second form, which I shall call indirect protection, includes two classes, the spiders which are specially protected themselves and those which mimic other creatures which are specially protected.

Spiders are specially protected when they become inedible through the acquisition of hard plates and sharp spines. The modification of form is frequently accompanied by conspicuous colors, which warn their enemies that they belong to an unpalatable class.

The second class of indirectly protected spiders--those that mimic specially protected creatures--presents some difficulties, since it is not always easy to determine whether the purpose of mimicry is protection or the capture of prey. The resemblance may, as is frequently the case in direct protection, serve both purposes.

In looking for instances of protective form and color among spiders we encounter one difficulty at the outset. The meaning of a protective peculiarity can be determined only when the animal is seen in its natural home. The number of strangely modified forms depicted in descriptive works on spiders is enormous. Bodies are twisted, elongated, inflated, flattened, truncated, covered with tubercles or spines, enclosed within chitinous plates, colored like bark, like lichens, like flowers of every imaginable hue, like bird droppings, like sand or stones, and in every one of these modifications there is doubtless an adaptation of the spider to its surroundings which, when it is studied out of its natural relations, we can only guess at.

It has been well said that in these protective resemblances those features of the portrait are most attended to by nature which produce the most effective deception when seen in nature; the faithfulness of the resemblance being much less striking when seen in the cabinet....

DIRECT PROTECTION. RESEMBLANCES TO VEGETABLE AND INORGANIC THINGS.--As a general rule the forms and colors of spiders are adapted to render them inconspicuous in their natural homes. Bright colored spiders, ... either keep hidden away or are found upon flowers whose tints harmonize with their own. This rule, while it has numerous exceptions, is borne out by the great majority of cases. A good illustration is found in the genus Uloborus, of which the members bear a deceptive resemblance to small pieces of bark, or to such bits of rubbish as commonly become entangled in old deserted webs. The only species in our neighborhood is Uloborus plumipes, which I have almost invariably found building in dead branches, where its disguise is more effective than it would be among fresh leaves. The spider is always found in the middle of the web, with its legs extended in a line with the body. There has been, in this species, a development along several lines, resulting in a disguise of considerable complexity. Its form and color make it like a scrap of bark, its body being truncated and diversified with small humps, while its first legs are very uneven, bearing heavy fringes of hair on the tibia and having the terminal joints slender. Its color is a soft wood-brown or gray, mottled with white. It has the habit of hanging motionless in the web for hours at a time, swaying in the wind like an inanimate object. The strands of its web are rough and inelastic, so that they are frequently broken; this gives it the appearance of one of those dilapidated and deserted webs in which bits of wind-blown rubbish are frequently entangled....

Out of seven examples of the species taken during one summer, five were found in dead tamarack branches, one on a dead bush, and the seventh, an interesting variety, under the eaves of a porch. My eye was caught by what seemed to be a string of eleven cocoons (it is not common to see more than four in a web). On attempting to take them down I was surprised to see one of the supposed cocoons begin to shake the web violently. Ten were what they seemed to be, but the eleventh was the mother spider, whose color and general appearance was exactly like that of the little cases that she had made for her eggs....

We come now to a large and interesting class in genus Epeira. I refer to those species, mostly nocturnal, which are protected during the day, not by hiding in crevices, nor in any way actually getting out of sight, but by the close resemblance which they bear to the bark of the trees to which they cling. This resemblance is brought about in two ways; through their color, which is like that of wood or lichens, and through their tuberculated and rugose forms, which resemble rough bark.

One of the most remarkable of these forms is C. mitralis, a Madagascar species, which, looked at in profile, probably resembles a woody knot. The abdomen is divided into two divergent cones (Fig. 1). The entire upper surface of the body is covered with conical elevations, which render it rough and uneven; the sides of the abdomen are made up of several layers, which form stages, one above another, like the ridges of bark on a woody excrescence. The legs, formed of wide, flattened plates, make the base. The color of the spider is yellowish-gray, varied with white and dark reddish-brown. It has the habit of perching on a branch and clasping it like a bird, so that the elaborate modification of form, which would be useless if the spider hung exposed in the web, is made as effective as possible.