Chapter VIII Leaf-Rolling Caterpillars
The labours of those insect-architects, which we have endeavoured to describe in the preceding pages, have been chiefly those of mothers to form a secure nest for their eggs, and the young hatched from them, during the first stage of their existence. But a much more numerous and not less ingenious class of architects may be found among the newly-hatched insects themselves, who, untaught by experience, and altogether unassisted by previous example, manifest the most marvellous skill in the construction of tents, houses, galleries, covert-ways, fortifications, and even cities, not to speak of subterranean caverns and subaqueous apartments, which no human art could rival.
The caterpillars, which are familiarly termed leaf-rollers, are perfect hermits. Each lives in a cell, which it begins to construct almost immediately after it is hatched; and the little structure is at once a house which protects the caterpillar from its enemies, and a store of food for its subsistence, while it remains shut up in its prison. But the insect only devours the inner folds. The art which these caterpillars exercise, although called into action but once, perhaps, in their lives, is perfect. They accomplish their purpose with a mechanical skill, which is remarkable for its simplicity and unerring success. The art of rolling leaves into a secure and immovable cell may not appear very difficult: nor would it be so if the caterpillars had fingers, or any parts which were equivalent to those delicate and admirable natural instruments with which man accomplishes his most elaborate works. And yet the human fingers could not roll a rocket-case of paper more regularly than the caterpillar rolls his house of leaves. A leaf is not a very easy substance to roll. In some trees it is very brittle. It has also a natural elasticity,—a disposition to spring back if it be bent,—which is caused by the continuity of its threads, or nervures. This elasticity is speedily overcome by the ingenuity with which the caterpillar works; and the leaf is thus retained in its artificial position for many weeks, under every variety of temperature. We will examine, in detail, how these little leaf-rollers accomplish their task.
One of the most common as well as the most simple fabrics constructed by caterpillars, may be discovered during summer on almost every kind of bush and tree. We shall take as examples those which are found on the lilac and on the oak.
A small but very pretty chocolate-coloured moth, abundant in every garden, but not readily seen, from its frequently alighting on the ground, which is so nearly of its own colour, deposits its eggs on the leaves of the currant, the lilac, and of some other trees, appropriating a leaf to each egg. As soon as the caterpillar is hatched, it begins to secure itself from birds and predatory insects by rolling up the lilac leaf into the form of a gallery, where it may feed in safety. We have repeatedly seen one of them when just escaped from the egg, and only a few lines long, fix several silk threads from one edge of a leaf to the other, or from the edge to the mid-rib; then going to the middle of the space, he shortened the threads by bending them with his feet, and consequently pulled the edges of the leaves into a circular form; and he retained them in that position by gluing down each thread as he shortened it. In their younger state, those caterpillars seldom roll more than a small portion of the leaf; but, when farther advanced, they unite the two edges together in their whole extent, with the exception of a small opening at one end, by which an exit may be made in case of need.
Another species of caterpillar, closely allied to this, rolls up the lilac leaves in a different form, beginning at the end of a leaf, and fixing and pulling its threads till it gets it nearly into the shape of a scroll of parchment. To retain this form more securely, it is not contented, like the former insect, with threads fixed on the inside of the leaf; but has also recourse to a few cables which it weaves on the outside.
Another species of moth, allied to the two preceding, is of a pretty green colour, and lays its eggs upon the leaves of the oak. This caterpillar folds them up in a similar manner, but with this difference, that it works on the under surface of the leaf, pulling the edge downwards and backwards, instead of forwards and upwards. This species is very abundant, and may readily be found as soon as the leaves expand. In June, when the perfect insect has appeared, by beating a branch of an oak, a whole shower of these pretty green moths may be shook into the air.
Among the leaf-rolling caterpillars, there is a small dark-brown one, with a black head and six feet, very common in gardens, on the currant-bush, or the leaves of the rose-tree (Lozotænia rosana, Stephens). It is exceedingly destructive to the flower-buds. The eggs are deposited in the summer, and probably also in the autumn or in spring, in little oval or circular patches of a green colour. The grub makes its appearance with the first opening of the leaves, of whose structure in the half-expanded state it takes advantage to construct its summer tent. It is not, like some of the other leaf-rollers, contented with a single leaf, but weaves together as many as there are in the bud where it may chance to have been hatched, binding their discs so firmly with silk, that all the force of the ascending sap, and the increasing growth of the leaves, cannot break through; a farther expansion is of course prevented. The little inhabitant in the meanwhile banquets securely on the partitions of its tent, eating door-ways from one apartment into another, through which it can escape in case of danger or disturbance.
The leaflets of the rose, it may be remarked, expand in nearly the same manner as a fan, and the operations of this ingenious little insect retain them in the form of a fan nearly shut. Sometimes, however, it is not contented with one bundle of leaflets, but by means of its silken cords unites all which spring from the same bud into a rain-proof canopy, under the protection of which it can feast on the flower-bud, and prevent it from ever blowing.
In the instance of the currant-leaves, the proceedings of the grub are the same; but it cannot unite the plaits so smoothly as in the case of the rose leaflets, and it requires more labour, also, as the nervures, being stiff, demand a greater effort to bend them. When all the exertions of the insect prove unavailing in its endeavours to draw the edges of a leaf together, it bends them inwards as far as it can, and weaves a close web of silk over the open space between. This is well exemplified in one of the commonest of our leaf-rolling caterpillars, which may be found as early as February on the leaves of the nettle and the white archangel (Lamium album). It is of a light dirty-green colour, spotted with black, and covered with a few hairs. In its young state it confines itself to the bosom of a small leaf, near the insertion of the leaf-stalk, partly bending the edges inwards, and covering in the interval with a silken curtain. As this sort of covering is not sufficient for concealment when the animal advances in growth, it abandons the base of the leaf for the middle, where it doubles up one side in a very secure and ingenious manner.
We have watched this little architect begin and finish his tent upon a nettle in our study, the whole operation taking more than half an hour. (J. R.) He began by walking over the plant in all directions, examining the leaves severally, as if to ascertain which was best fitted for his purpose by being pliable, and bending with the weight of his body. Having found one to his mind, he placed himself along the mid-rib, to the edge of which he secured himself firmly with the pro-legs of his tail; then stretching his head to the edge of the leaf, he fixed a series of parallel cables between it and the mid-rib, with another series crossing these at an acute angle. The position in which he worked was most remarkable, for he did not, as might have been supposed, spin his cables with his face to the leaf, but throwing himself on his back, which was turned towards the leaf, he hung with his whole weight by his first-made cables. This, by drawing them into the form of a curve, shortened them, and consequently pulled the edge of the leaf down towards the mid-rib. The weight of his body was not, however, the only power which he employed; for, using the terminal pro-legs as a point of support, he exerted the whole muscles of his body to shorten his threads, and pull down the edge of the leaf. When he had drawn the threads as tight as he could, he held them till he spun fresh ones of sufficient strength to retain the leaf in the bent position into which he had pulled it. He then left the first series to hang loose while he shortened the fresh-spun ones as before. This process was continued till he had worked down about an inch and a half of the leaf, as much as he deemed sufficient for his habitation. This was the first part of the architecture.
By the time he had worked to the end of the fold, he had brought the edge of the leaf to touch the mid-rib; but it was only held in this position by a few of the last-spun threads, for all the first-spun ones hung loose within. Apparently aware of this, the insect protruded more than half of its body through the small aperture left at the end, and spun several bundles of threads on the outside precisely similar to those ropes of a tent which extend beyond the canvas, and are pegged into the ground. Unwilling to trust the exposure of his whole body on the outside, lest he should be seized by the first sand-wasp (odynerus) or sparrow which might descry him, he now withdrew to complete the internal portion of his dwelling, where the threads were hanging loose and disorderly. For this purpose he turned his head about, and proceeded precisely as he had done at the beginning of his task, but taking care to spin his new threads so as to leave the loose ones on the outside, and make his apartment smooth and neat. When he again reached the opposite end, he constructed there also a similar series of cables on the outside, and then withdrew to give some final touches to the interior.
It is said by Kirby and Spence,BJ that when these leaf-rolling insects find that the larger nervures of the leaves are so strong as to prevent them from bending, they “weaken it by gnawing it here and there half through.” We have never observed the circumstance, though we have witnessed the process in some hundreds of instances; and we doubt the statement, from the careful survey which the insect makes of the capabilities of the leaf before the operation is begun. If she found upon examination that a leaf would not bend, she would reject it, as we have often seen happen, and pass to another. (J. R.)
A species of leaf-roller, of the most diminutive size, merits particular mention, although it is not remarkable in colour or figure. It is without hair, of a greenish-white, and has all the vivacity of the other leaf-rollers. Sorrel is the plant on which it feeds; and the manner in which it rolls a portion of the leaf is very ingenious.
The structure which it contrives is a sort of conical pyramid, composed of five or six folds lapped round each other. From the position of this little cone the caterpillar has other labours to perform, beside that of rolling the leaf. It first cuts across the leaf, its teeth acting as a pair of scissors; but it does not entirely detach this segment. It rolls it up very gradually, by attaching threads of silk to the plane surface of the leaf, as we have before seen; and then, having cut in a different direction, sets the cone upright, by weaving other threads, attached to the centre of the roll and the plane of the leaf, upon which it throws the weight of its body. This, it will be readily seen, is a somewhat complicated effort of mechanical skill. It has been minutely described by M. Réaumur; but the preceding representation will perhaps make the process clearer than a more detailed account.
This caterpillar, like those of which we have already spoken, devours all the interior of the roll. It weaves, also, in the interior, a small and thin cocoon of white silk, the tissue of which is made compact and close. It is then transformed into a chrysalis.
The caterpillars of two of our largest and handsomest butterflies, the painted lady (Cynthia cardui, Stephens), and the admiral, or Alderman of the London fly-fanciers (Vanessa atalanta), are also leaf-rollers. The first selects the leaves of the great spear-thistle, and sometimes those of the stemless or star-thistle, which might be supposed rather difficult to bend; but the caterpillar is four times as large and strong as those which we have been hitherto describing. In some seasons it is plentiful; in others it is rarely to be met with: but the admiral is seldom scarce in any part of the country; and by examining the leaves of nettles which appear folded edge to edge, in July and August, the caterpillar may be readily found.
Another butterfly (Hesperia malvæ) is met with on dry banks where mallows grow, in May, or even earlier, and also in August, but is not indigenous. The caterpillar, which is grey, with a black head, and four sulphur-coloured spots on the neck, folds around it the leaves of the mallow, upon which it feeds. There is nothing, however, peculiarly different in its proceedings from those above described; but the care with which it selects and rolls up one of the smaller leaves, when it is about to be transformed into a chrysalis, is worthy of remark; it joins it, indeed, so completely round and round, that it has somewhat the resemblance of an egg. Within this green cell it lies secure, till the time arrives when it is ready to burst its cerements, and trust to the quickness of its wings for protection against its enemies.
Among the nests of caterpillars which roll up parcels of leaves, we know none so well contrived as those which are found upon willows and a species of osier. The long and narrow leaves of these plants are naturally adapted to be adjusted parallel to each other; for this is the direction which they have at the end of each stalk, when they are not entirely developed. One kind of small smooth caterpillar (Tortrix chlorana), with sixteen feet, the under part of which is brown, and streaked with white, fastens these leaves together, and makes them up into parcels. There is nothing particularly striking in the mechanical manner in which it constructs them. It does precisely what we should do in a similar case: it winds a thread round those leaves which must be kept together, from a little above their termination to a very short distance from their extreme point; and as it finds the leaves almost constantly lying near each other, it has little difficulty in bringing them together, as is shown in the following cut, a.
The prettiest of these parcels are those which are made upon a kind of osier, the borders of whose leaves sometimes form columnar bundles before they become developed. A section of these leaves has the appearance of filigree-work (see b).
A caterpillar which feeds upon the willow, and whose singular attitudes have obtained for it the trivial name of Ziczac, also constructs for itself an arbour of the leaves, by drawing them together in an ingenious manner. M. RoeselBK has given a tolerable representation of this nest, and of the caterpillar. The caterpillar is found in June; and the moth (Notodonta ziczac) from May to July in the following year (see cut, p. 151).
Beside those caterpillars which live solitary in the folds of a leaf, there are others which associate, employing their united powers to draw the leaves of the plants they feed upon into a covering for their common protection. Among these we may mention the caterpillar of a small butterfly, the plantain or Glanville fritillary (Melitea cinxia), which is very scarce in this country.
Although a colony of these caterpillars is not numerous, seldom amounting to a hundred individuals, the place which they have selected is not hard to discover. Their abode may be seen in the meadow in form of a tuft of herbage covered with a white web, which may readily be mistaken, at first view, for that of a spider, but closer inspection soon corrects this notion. It is, in fact, a sort of common tent, in which the whole brood lives, eats, and undergoes the usual transformations. The shape of this tent, for the most part, approaches the pyramidal, though that depends much upon the natural growth of the herbage which composes it. The interior is divided into compartments formed by the union of several small tents, as it were, to which others have been from time to time added according to the necessities of the community.
When they have devoured all the leaves, or at least those which are most tender and succulent, they abandon their first camp, and construct another contiguous to it under a tuft of fresh leaves. Several of these encampments may sometimes be seen within the distance of a foot or two, when they can find plantain (Plantago lanceolata) fit for their purpose; but though they prefer this plant, they content themselves with grass if it is not to be procured.
When they are about to cast their skins, but particularly when they perceive the approach of winter, they construct a more durable apartment in the interior of their principal tent. The ordinary web is thin and semi-transparent, permitting the leaves to be seen through it; but their winter canvas, if we may call it so, is thick, strong, and quite opaque, forming a sort of circular hall without any partition, where the whole community lie coiled up and huddled together.
Early in spring they issue forth in search of fresh food, and again construct tents to protect them from cold and rain, and from the mid-day sun.
M. Réaumur found upon trial, that it was not only the caterpillars hatched from the eggs of the same mother which would unite in constructing the common tent; for different broods, when put together, worked in the same social and harmonious manner. We ourselves ascertained, during the present summer (1829), that this principle of sociality is not confined to the same species, nor even to the same genus. The experiment which we tried was to confine two broods of different species to the same branch, by placing it in a glass of water to prevent their escape. The caterpillars which we experimented on were several broods of the brown-tail moth (Porthesia auriflua) and the lackey (Clisiocampa neustria). These we found to work with as much industry and harmony in constructing the common tent as if they had been at liberty on their native trees; and when the lackeys encountered the brown-tails they manifested no alarm nor uneasiness, but passed over the backs of one another, as if they had made only a portion of the branch. In none of their operations did they seem to be subject to any discipline, each individual appearing to work, in perfecting the structure, from individual instinct, in the same manner as was remarked by M. Huber in the case of the hive-bees. In making such experiments, it is obvious that the species of caterpillars experimented with must feed upon the same sort of plant.BL (J. R.)
The design of the caterpillars in rolling up the leaves is not only to conceal themselves from birds and predatory insects, but also to protect themselves from the cuckoo-flies, which lie in wait in every quarter to deposit their eggs in their bodies, that their progeny may devour them. Their mode of concealment, however, though it appear to be cunningly contrived and skilfully executed, is not always successful, their enemies often discovering their hiding-place. We happened to see a remarkable instance of this last summer (1828), in the case of one of the lilac caterpillars which had changed into a chrysalis within the closely-folded leaf. A small ichneumon, aware it should seem of the very spot where the chrysalis lay within this leaf, was seen boring through it with her ovipositor, and introducing her eggs through the punctures thus made into the body of the dormant insect. We allowed her to lay all her eggs, about six in number, and then put the leaf under an inverted glass. In a few days the eggs of the cuckoo-fly were hatched, the grubs devoured the lilac chrysalis, and finally changed into pupæ in a case of yellow silk, and into perfect insects like their parents. (J. R.)