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[Ill.u.s.tration: FIG. 1. THE CATERPILLAR OF THE MARBLED WHITE b.u.t.tERFLY (_Arge galathea_).]
In fact, whenever in any group we find differences in form or color, we shall always find them a.s.sociated 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 gra.s.ses), 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 gra.s.ses.
[Ill.u.s.tration: FIG. 2.--THE CATERPILLAR OF THE EYED HAWK-MOTH (_Smerinthus ocellatus_).]
Now, similar lines occur on a great number of caterpillars belonging to most different groups of b.u.t.terflies and moths, as you may see by turning over the ill.u.s.trations 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 b.u.t.terflies, especially in those which feed on gra.s.s; 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 gra.s.s. 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 b.u.t.terflies 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 gra.s.s; 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 att.i.tudes, 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 b.u.t.terfly also has whitish caterpillars, and this may at first sight appear to contradict the rule, since it feeds on gra.s.s. 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 princ.i.p.al categories: firstly, those which, like the Geometridae, put themselves into peculiar and stiff att.i.tudes, 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 b.u.t.terfly, 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 b.u.t.terflies or moths which feed on the pine, but there are a few: and most, if not all of them, have a very a.n.a.logous 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.
[Ill.u.s.tration: FIG. 3.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). Full grown. Natural size.]
[Ill.u.s.tration: FIG. 4.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). First stage.]
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.
[Ill.u.s.tration: FIG. 5.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). Second Stage.]
[Ill.u.s.tration: FIG. 6.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). Just before the second moult.]
[Ill.u.s.tration: FIG. 7.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). Third Stage.]
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 a.s.sume 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.
[Ill.u.s.tration: FIG. 8.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). Fourth Stage.]
Now, the princ.i.p.al 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.
[Ill.u.s.tration: FIG. 9.--THE CATERPILLAR OF THE ELEPHANT HAWK-MOTH (_Chaerocampa elpenor_). Fifth Stage.]
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 circ.u.mstances 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 att.i.tudes, 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 const.i.tute 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 a.s.sembled 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.
[Ill.u.s.tration: FIG. 10.--THE CATERPILLAR OF THE SMALL ELEPHANT HAWK-MOTH (_Chaerocampa porcellus_).]
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 pa.s.ses through in a few weeks. Thus, the caterpillar of _Chaerocampa porcellus_, a species very nearly allied to the Elephant Hawk-moth, pa.s.ses 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 pa.s.sed 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 a.s.sumed. 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 b.u.t.terflies 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 b.u.t.terflies and moths are as easy to distinguish as the perfect insects; they differ from one another as much as, sometimes more than, the b.u.t.terflies and moths themselves.
There are five princ.i.p.al 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 a.s.sume 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 att.i.tudes, 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 gra.s.s 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 const.i.tuting 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 gra.s.s 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 b.u.t.terfly is not very conspicuous, and has been omitted in some descriptions. Blue is even rarer than red; in fact, none of our b.u.t.terfly 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 att.i.tudes they a.s.sume. 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 att.i.tudes.