The Life-Story of Insects

by George H. Carpenter

Most remarkable, in many respects, of all aquatic larvae are the grubs of the Sand-midges (Simulium). These live entirely submerged and, having no special gills, carry out an exchange of gases through the general surface of the cuticle between the dissolved air in the water and the cavities of the air-tube system. The body is shaped like a flask swollen slightly at the hinder end and possesses a median pro-leg just behind the head, also another at the tail, which serves to attach the larva to a stone or to the leaf of an aquatic plant. The head has, in addition to feelers and jaws, a pair of processes with wonderful fringes which by their motion set up currents in the water, and bring food particles within reach of the mouth. A number of the larvae usually live in a community. Their power of spinning silken threads by which they can work their way back when accidentally dislodged from their resting-place, has been vividly described by [Miall (1895)].

Examples might be multiplied, but enough have been given to enforce the conclusion that the forms of insect-larvae are wondrously varied, and that frequently, within the limits of the same order or even family, modifications of type may be found which are suited to various modes of life adopted by different insects. A survey of the multitudes of insect larvae—grubs, caterpillars, maggots—living on land, on plants, underground, in the water; feeding on leaves, in stems, on roots, on carrion, on refuse; by hunting or by lurking after prey; as parasites or as scavengers, brings home to us most strongly the conclusion that each larva is fitted to some little niche in the vast temple of life, each is specially adapted to its part in the great drama of being.

CHAPTER VII
PUPAE AND THEIR MODIFICATIONS

The pupal stage is characteristic of the life-story of those insects whose larvae have wing-rudiments in the form of inpushed imaginal discs, and in all these insects there is, as we have seen, considerable divergence in form between larva and imago. In the pupa the wings and other characteristically adult structures are, for the first time, visible outwardly; it is the instar which marks the great crisis in transformation. The pupa rests, as a rule, in a quiescent condition, and during the early period of this stage the needful internal changes, the breaking down of many larval tissues, and their replacement by imaginal organs, go on. Both outwardly and inwardly therefore, the insect undergoes, at the pupal stage, a reconstruction necessitated by the differences in form and often in habit, between the larva and the winged adult; and the greater these differences, the more profound must be the changes that mark the pupal stage.

From the prominence of imaginal structures in the pupa, it is at once seen that the pupa of any insect must resemble the adult more nearly than it resembles the larva. But in different groups of insects we find different degrees of likeness between pupa and imago. In a beetle pupa (see [fig. 16] c), the appendages—feelers, jaws, legs, wings—stand out from the body as do those of the perfect insect. This type is called a free pupa. The pupal cuticle has to be shed for the emergence of the imago, but the pupa is already a somewhat reduced model of the final instar, with abbreviated wings and doubled-up legs. A free pupa is characteristic of the Coleoptera, Neuroptera, Trichoptera, Hymenoptera and many Diptera. In some cases the pupa requires to be specially adapted for a peculiar mode of life; for example, a special arrangement of breathing organs may be necessary for life under water, and there must needs be temporary pupal structures, not represented in the imago.

On the other hand, in the pupae of most Lepidoptera and of some Diptera, there is more or less coalescence between the cuticle of the appendages and the cuticle of the body generally, so that the appendages do not stand out, but being, as it were, glued down to the body, are somewhat masked (see [fig. 1] e and [fig. 23]). Consequently the obtect pupa, as this type is called, does not resemble its imago as fully as a free pupa does. The outline of the wings for example in a butterfly's pupa can in some cases be traced only with difficulty. T. A. Chapman has shown [(1893)] that the completely obtect pupa characterises the more highly developed families of Lepidoptera, while in the more primitive families the pupa is incompletely obtect. If the pupa of a butterfly or moth be lifted and held in the hand, a bending or wriggling motion of the abdomen can be observed. In the incompletely obtect pupa, this motion is evident in a greater number of segments than in the completely obtect, the number concerned varying from five to two in different families. In the nymphalid butterflies, the pupa is often called a 'chrysalis' on account of the golden hue displayed by the cuticle, and the term 'chrysalis' is sometimes bestowed indiscriminately on any kind of pupa. It has been shown by [Poulton (1892)] and others, that the colour of a butterfly pupa is to some extent affected by the surroundings of the caterpillar just before its last moult.

Reference has been made ([p. 58]) to the power of spinning silk possessed by many larvae; often the principal use of this silk is to form some protection for the pupa, the larva before its last moult constructing a cocoon within which the pupa may rest safely. Many larvae bury themselves in the earth, and the pupa lies in an earthen chamber, the lining particles of soil fastened together by fine silken threads. Larvae that feed in wood, like the caterpillar of the Goat-moth (Cossus) make a cocoon of splinters spun together, while hairy caterpillars, such as those of the Tiger-moths, work some of their hairs in with the silk to make a firm cocoon ([fig. 17] b). On the other hand, those caterpillars known as 'silkworms' make a dense cocoon of pure silk, consisting of two layers, the outer of coarse and the inner of fine threads. Silken cocoons very similar in appearance are spun by the larvae of small Ichneumon-flies. Many pupae lie in a loose cocoon formed of a few interlacing threads, as for example the conspicuous black and yellow banded pupa of the Magpie-moth (Abraxas grossulariata) and the pupae of various leaf-beetles. Others again spin together the edges of leaves with connecting silken threads. The grubs of bees and wasps which are reared in the comb-chambers of their nests seal up the opening of the chamber with a lid, partly silk ([fig. 18] co) and partly excretion, when ready to pass into the pupal state. An additional external 'capping' may be also supplied by the workers.

The pupae of butterflies are especially interesting, as illustrating the extreme reduction of the silken cocoon. The pupa of a 'swallowtail' (Papilionid) or a 'white' (Pierid) butterfly ([fig. 23]) may be found attached to a twig of its food-plant or to a wall, in an upright position, its tail fastened to a pad of silk and a slender silken girdle encircling its thorax. The pupa of a 'Tortoiseshell' or 'Admiral' (Nymphalid) butterfly hangs head downwards from a twig, supported only by the tail-pad of silk, which, useless as a shelter, serves only for attachment. The pupa is fastened to this pad by a spiny hook or process, the cremaster ([fig. 23] cr), on the last abdominal segment. The cremaster is a characteristic structure in the pupa of a moth or butterfly. [C. V. Riley (1880)] and [W. Hatchett-Jackson (1890)] have shown that it corresponds with a spiny area, the suranal plate, which lies above the opening of the caterpillar's intestine. The means by which the suspended pupa of a nymphalid butterfly attaches its cremaster to the silken pad which the larva has spun in preparation for pupation, is worthy of brief attention. The caterpillar, hanging head downwards, is attached to the silken pad by its hindmost pair of pro-legs or claspers and by the suranal plate, and the cuticle is slowly worked off from before backwards, so as to expose the pupa. Were the process of moulting to be simply completed while the insect hangs by the claspers, the pupa would of course fall to the ground. But there is enough adhesion between the pupal and larval cuticles at the hinder end of the body, especially by means of the everted lining of the hind-gut, for the pupa to be supported while it jerks its cremaster out of the larval cuticle and works it into the meshes of the silken pad. The moult is thus completed and the pupa hangs securely all the time. In the numerous cases where the pupa is enclosed in a cocoon, the cremaster serves to fix the pupa to the surrounding silk. [Chapman (1893)] has drawn attention to the fact that among the more highly organised moths the pupa remains in the cocoon, the emergence being entirely left to the imago, while the pupae of the more primitive moths work their way partly out of the cocoon before the final moult begins. In the latter case, the cremaster is anchored by a strand of silk which allows a certain degree of emergence, and the pupa has rows of spines on its abdominal segments, of which a greater number retain the power of mutual motion than in those pupae which do not come out of their cocoons.