Part 23

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.

[Illustration: 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 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.

[Illustration]

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.

[Illustration]

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.

[Illustration: FIG. 1.--CAEROSTRIS MITRALIS (from Vinson).]

[Illustration: FIG. 2.--CAEROSTRIS MITRALIS, in profile (from Vinson).]

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.