The Life-Story of Insects. George H. Carpenter

[3] The term 'hypodermis' frequently applied to this layer is misleading. The layer is the true outer skin—ectoderm or epidermis.

      Now this jointed exoskeleton—an admirably formed suit of armour though it often is—has one drawback: it is not part of the insect's living tissues. It is a cuticle formed by the solidifying of a fluid secreted by the epidermal cells, therefore without life, without the power of growth, and with only a limited capacity for stretching. It follows, therefore, that at least during the period through which the insect continues to grow, the cuticle must be periodically shed. Thus in the life-story of an insect or other arthropod, such as a lobster, a spider, or a centipede, there must be a succession of cuticle-castings—'moults' or ecdyses as they are often called.

      When such a moult is about to take place the cuticle separates from the underlying epidermis, and a fluid collects beneath. A delicate new cuticle (see fig. 10 cu') is then formed in contact with the epidermis, and the old cuticle opens, usually with a slit lengthwise along the back, to allow the insect in its new coat to emerge. At first this new coat is thin and flabby, but after a period of exposure to the air it hardens and darkens, becoming a worthy and larger successor to that which has been cast. The cuticle moreover is by no means wholly external. The greater part of the digestive canal and the whole air-tube system are formed by inpushings of the outer skin (ectoderm) and are consequently lined with an extension of the chitinous cuticle which is shed and renewed at every moult.

      In all insects these successive moults tend to be associated with change of form, sometimes slight, sometimes very great. The new cuticle is rarely an exact reproduction of the old one, it exhibits some new features, which are often indications of the insect's approach towards maturity. Even in some of those interesting and primitive insects the Bristle-tails (Thysanura) and Spring-tails (Collembola), in which wings are never developed, perceptible differences in the form and arrangement of the abdominal limbs can be traced through the successive stages, as R. Heymons (1906) and K. W. Verhoeff (1911) have shown for Machilis. But the changes undergone by such insects are comparatively so slight, that the creatures are often known as 'Ametabola' or insects without transformation in the life-history. Now there are a considerable number of winged insects—cockroaches and grasshoppers for example—in which the observable changes are also comparatively slight. We will sketch briefly the main features of the life-story of such an insect.

      Fig. 4. Common Cockroach (Blatta orientalis). a, female; b, male; c, side view of female; d, young. After Marlatt, Entom. Bull. 4, U.S. Dept. Agric.

The young creature is hatched from the egg in a form closely resembling, on the whole, that of its parent, so that the term 'miniature adult' sometimes applied to it, is not inappropriate. The baby cockroach (fig. 4 d) is known by its flattened body, rounded prothorax, and stiff, jointed tail-feelers or cercopods; the baby grasshopper by its strong, elongate hind-legs, adapted, like those of the adult, for vigorous leaping. During the growth of the insect to the adult state there may be four or five moults, each preceded and succeeded by a characteristic instar[4]. The first instar differs, however, from the adult in one conspicuous and noteworthy feature, it possesses no trace of wings. But after the first or the second moult, definite wing-rudiments are visible in the form of outgrowths on the corners of the second and third thoracic segments. In each succeeding instar these rudiments become more prominent, and in the fourth or the fifth stage, they show a branching arrangement of air-tubes, prefiguring the nervures of the adult's wing (fig. 5). After the last moult the wings are exposed, articulated to the segments that bear them, and capable of motion. Having been formed beneath the cuticle of the wing-rudiments of the penultimate instar, the wings are necessarily abbreviated and crumpled. But during the process of hardening of the cuticle, they rapidly increase in size, blood and air being forced through the nervures, so that the wings attaining their full expanse and firmness, become suited for the function of flight.

[4] The convenient term 'instar' has been proposed by Fischer and advocated by Sharp (1895) for the form assumed by an insect during a stage of its life-story. Thus the creature as hatched from the egg is the first instar, after the first moult it has become the second instar, and so on, the number of moults being always one less than the number of instars.

      Fig. 5. Nymph of Locust (Schistocera americana) with distinct wing-rudiments. After Howard, Insect Life, vol. VII.

      The changes through which these insects pass are therefore largely connected with the development of the wings. It is noteworthy that in an immature cockroach the entire dorsal cuticle is hard and firm. In the adult, however, while the cuticle of the prothorax remains firm, that of the two hinder thoracic and of all the abdominal segments is somewhat thin and delicate on the dorsal aspect. It needs not now to be resistant, because it is covered by the two firm forewings, which shield and protect it, except when the insect is flying. There are, indeed, slight changes in other structures not directly connected with the wings. In a young grasshopper, for example, the feelers are relatively stouter than in the adult, and the prothorax does not show the specifically distinctive shape with its definite keels and furrows. Changes in the secondary sexual characters may also be noticed. For instance, in an immature cockroach both male and female carry a pair of jointed tail-feelers or cercopods on the tenth abdominal segment, and a pair of unjointed limbs or stylets on the ninth. In the adult stage, both sexes possess cercopods, but the males only have stylets, those of the female disappearing at the final moult.

      Reviewing the main features of the life-story of a grasshopper or cockroach, we notice that there is no marked or sudden change of form. The newly-hatched insect resembles generally its parent, except that it has no wings. Wing-rudiments appear, however, in an early instar as visible outgrowths on the thoracic segments, and become larger after each moult. All through its various stages the immature insect—nymph as it is called—lives in the same kind of situations and on the same kind of food as its parent, and it is all along active and lively, undergoing no resting period like the pupal stage in the transformation of the butterfly.

      One interesting and suggestive fact remains to be mentioned. There are grasshoppers and cockroaches in which the changes are even less than those just sketched, because the wings remain, even in the adult, in a rudimentary state (as for example in the female of the common kitchen cockroach, Blatta orientalis, see fig. 4 a), or are never developed at all. Such exceptional winglessness in members of a winged family can only be explained by the recognition of a life-story, not merely in the individual but in the race. We cannot doubt that the ancestors of these wingless insects possessed wings, which in the course of time have been lost by the whole species or by the members of the female sex. It is generally assumed that this loss has been gradual, and so in many cases it probably may have been. But there are species of insects in which some generations are winged and others wingless; a winged mother gives birth to wingless offspring, and a wingless parent to young with well-developed wings. Such discontinuity in the life-story of a single generation forces us to recognise the possibility of similar sudden mutations in the course of that age-long process of evolution to which the facts of insect growth, and indeed of all animal development, bear striking testimony.

CHAPTER III

       THE LIFE-STORIES OF SOME SUCKING INSECTS

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