George H. Carpenter

“ Perhaps the most striking result of the study of insect transformation is the appreciation of the divergent specialisation of larva and imago, and it is a suggestive thought that of the two the larva has in many cases diverged the more from the typical condition. The caterpillar crawling over the leaf, or the fly-grub swimming through the water, may thus be regarded as a creature preparing for a change to the true conditions of its life. It is a strange irony that the preparation is often far longer than the brief hours of achievement. ”

“ Our study of the life-story of insects, therefore, while it has taught us something of what is going on around us to-day, and has given us hints of the course of a few threads of that long life-story which runs through the ages, brings us face to face with the most instructive, if humbling fact that 'there are many more things of which we are ignorant.' The passage from creeping to flight, as the caterpillar becomes transformed into the butterfly, was a mystery to those who first observed it, and many of its aspects remain mysterious still. ”

“ 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. ”

“ The fly—with its large globular head, bearing the extensive compound eyes, the highly modified feelers with their exquisitely feathered slender sensory tips, and the complex suctorial jaws; with its compact thorax bearing the glassy fore-wings alone used for flight, though the hind-wings modified into tiny drumstick-like 'halters' are the organs of a fine equilibrating sense—is perhaps the most specialised, structurally the 'highest' of all insects. ”

“ The aquatic young of a stone-fly does not differ sufficiently in form from its parent to warrant us in calling it a larva; the life-history is like that of a cockroach, all the instars however except the final one—the winged adult or imago—live in the water. The young of one of our large species, a Perla for example, has well-chitinised cuticle, broad head, powerful legs, long feelers and cerci like those of the imago; its wings arise from external rudiments, which are conspicuous in the later aquatic stages. ”

“ The foregoing brief outline of our knowledge of the geological succession of insects shows that the exopterygote preceded, in time, the endopterygote type of life-history. We have already seen that those insects undergoing little change in the life-cycle, and with visible, external wing-rudiments, are on the whole less specialised in structure than those which pass through a complete transformation. These two considerations, taken together, suggest strongly that in the evolution of the insect class, the simpler life-history preceded the more complex. ”

“ And it is of interest to note that this somewhat degraded type of larva is remarkably constant through a great series of families—gall-flies, ichneumon-flies, wasps, bees (fig. 18) , ants—that vary widely in the details of their structure and in their habits and mode of life. Almost without exception, however, they make in some way abundant provision for their young. The feeble, helpless, larva is in every case well sheltered and well fed; it has not to make its own way in the world, as the active armoured larva of a ground-beetle or the caterpillar of a butterfly is obliged to do. ”

“ This is not surprising when we remember that there is often a great difference between larva and imago in the nature of the food. The digestive canal of a caterpillar runs a fairly straight course through the body and consists of a gullet, stomach (mid-gut) , intestine, and rectum; it is adapted for the digestion of solid food. In the butterfly there is one outgrowth of the gullet in the head—a pharyngeal sac adapted for sucking liquids; and another outgrowth at the hinder end of the gullet (which is much longer than in the larva) —a crop or food-reservoir lying in the abdomen. ”

“ During many centuries naturalists have bent their energies to explain the difficulties presented by insect transformations. Aristotle, the first serious student of organised beings whose writings have been preserved for us, and William Harvey, the famous demonstrator of the mammalian blood circulation two thousand years later, agreed in regarding the pupa as a second egg. The egg laid by a butterfly had not, according to Harvey, enough store of food to provide for the building-up of a complex organism like the parent; only the imperfect larva could be produced from it. ”

“ The vast majority of caterpillars feed on the leaves of plants and their long worm-like bodies with the series of paired pro-legs, are excellently adapted for their habit of clinging to twigs, and crawling along shoots or the edges of leaves as they go in search of food. Of great importance to a caterpillar is its power of spinning silk, consisting of fine threads solidified from the secretion of specially modified salivary glands whose ducts open in the insect's mouth at the tip of the tubular tongue which forms a spinneret. ”

“ 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. ”

“ The caterpillar differs markedly from the butterfly. As we pursue our studies of insect growth and transformation we shall find that in some cases the difference between young and adult is much greater—as for example between the maggot and the house-fly, in others far less—as between the young and full-grown grasshopper or plant-bug. It is evidently wise to begin a general survey of the subject with some of those simpler cases in which the differences between the young and adult insect are comparatively slight. ”

“ The female loses legs and feelers, and never acquires wings, becoming little more than a sluggish egg-bag (fig. 7 e) . The male on the other hand passes into a second larval stage in which there are no functional legs, but rudiments of legs and of wings are present on the epidermis beneath the cuticle, as shown by B. O. Schmidt for Aspidiotus (1885) . The penultimate instar of this sex in which the wing-rudiments are visible externally lies passively beneath the scale, its behaviour resembling that of a butterfly pupa. The adult winged male (fig. 7 a) leads a short, but active life. ”

“ It is noteworthy that in these short-lived insects the male is often provided with elaborate sense-organs which, we may believe, assist him to find a mate with as little delay as possible; the male may-fly has especially complex eyes, while the feelers of the male silk-moth or eggar are comb-like or feathery, the branches bearing thousands of sensory hairs. A box with a captive living female of one of these moths, if taken into a wood haunted by the species becomes rapidly surrounded by a swarm of would-be suitors, attracted by the odour emitted from the prisoner's scent-glands. ”

“ Here we see that the feeding activities of the insect are carried on during the larval stage only; the may-fly in its winged condition takes no food, pairing and egg-laying form the whole of its appointed task. A similar though less extreme shortening of the imaginal life may be noticed in many endopterygote insects. For example, the bot- and warble-flies have the jaws so far reduced that they are unable to feed, and the parasitic life of the maggot (see p. 74) extending over eight or nine months in the body of the horse or ox, prepares for a winged existence of probably but a few days. ”

“ A large part of the substance of an insect larva consists of muscular tissue, surrounding the digestive tube, and forming the great muscles that move the various parts of the body, and of fat, surrounding the organs and serving as a store of food-material. Very many of the muscle-fibres and the fat-cells also become disintegrated during the late larval and pupal stages, and the corresponding tissues of the adult are new formations derived from special groups of imaginal cells, though some muscles may persist from the larva to the adult. ”

“ The life-story of the butterfly, then, well-known as it may be, furnishes a puzzling exception to some wide-reaching generalisations concerning animal development. And the student of science often finds that an exception to some rule is the key to a problem of the highest interest. ”

“ 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 ”

“ At the same time of course, owing to the extended term of the larval life, many more individuals of the species are wintering underground as 'wireworms' of various ages, and these, except in very severe frosts, can continue their occupation of feeding on roots. But in the case of the 'turnip-flies' the food-supply is cut off in winter, and all those beetles of the latest summer brood that survive hibernate in some sheltered spot, waiting for the return of spring, that they may lay their eggs, and start the life-cycle once again. ”

“ Everyone is familiar with the main facts of such a life-story as that of a moth or butterfly. The form of the adult insect (fig. 1 a) is dominated by the wings—two pairs of scaly wings, carried respectively on the middle and hindmost of the three segments that make up the thorax or central region of the insect's body. Each of these three segments carries a pair of legs. In front of the thorax is the head on which the pair of long jointed feelers and the pair of large, sub-globular, compound eyes are the most prominent features. ”

“ An insect that is continually submerged and has no contact with the upper air cannot breathe through a series of paired spiracles, and during the aquatic life-period of the stone-fly these remain closed. Nevertheless, breathing is carried on by means of the ordinary system of branching air-tubes, the trunks of which are in connection with the tufted hollow gill-filaments, through whose delicate cuticle gaseous exchange can take place, though the method of this exchange is as yet very imperfectly understood. ”

“ Whatever differences of opinion may prevail on points of detail, the general explanation of insect metamorphosis as the result of divergent evolution in the two active stages of the life-story must assuredly be accepted. No other explanation accords with the increasing degree of divergence to be observed as we pass from the lower to the higher insect orders. ”

“ A butterfly pupa, like the imago, has no mandibles, but in the case of the Caddis-flies (Trichoptera) and two families of small moths, the most primitive of all Lepidoptera, the pupa, like the larva, has well-developed mandibles. These enable the caddis pupa to bite its way out of the shortened larval case in which it has pupated, and then to swim upwards through the water ready for the caddis-fly's emergence into the air. Pupae that are submerged require special breathing-organs. ”

“ After a long subterranean existence, usually extending over several years, the insect attains the penultimate stage of its life-story, during which it rests passively within an earthen cell, awaiting the final moult, which will usher in its winged and perfect state. ”

“ The dragon-fly larva has the same predaceous mode of life as its parent, but it is sluggish in habit, lurking for its prey at the bottom of the pond, among the mud or vegetation, which it resembles in colour. ”

“ Reviewing the small selection of life-stories of various Lepidoptera just sketched, we notice an interesting and suggestive variety in the wintering stage. The vanessid butterflies hibernate as imagos; the 'vapourer' winters in the egg, the magpie as a young ungrown larva, the 'tiger' as a half-size larva; the Agrotis caterpillar feeds through the winter, growing all the time; the codling caterpillar completes its growth in the autumn, and winters as a full-size resting larva ”

“ Small insects have, as a rule, shorter lives than large ones; those whose larvae devour highly nutritive food generally develop more quickly than those which have to live on dry, poor, substances; life-cycles follow one another most rapidly in summer weather when temperature is high and food plentiful.
In early chapters we have already noticed the long aquatic life of the larva and nymph of a dragon-fly, relatively a large insect, and the rapid multiplication of the repeated summer broods of virgin aphids (p. 18) . ”

“ Such division of the life-history into a long feeding, and a short breeding period has, as will be seen later, an important bearing on the question of insect transformation generally, and the dragon-flies and may-flies afford examples of two stages in its specialisation. The sub-imaginal instar of the may-fly furnishes also a noteworthy fact for comparison with other insect histories. In two points, however, the life-story of these flies with their aquatic larvae recalls that of the cockroach. ”

“ The grubs of the Chafers (Scarabaeidae) are also root-eaters, but they are less active in their habits than the wireworms, and the cuticle of their somewhat stout bodies is, for the most part, pale and flexible; only the head and legs are hard and horny. Usually an evident correspondence can be traced between the outward form of any larva and its mode of life. For example, in the family of the Leaf-beetles (Chrysomelidae) some larvae feed openly on the foliage of trees or herbs, while others burrow into the plant tissues. ”

“ We have just seen that a caterpillar is the usual form of larva among the saw-flies, but in all other families of the Hymenoptera we find the legless grub. A grub of this order may usually be distinguished from the larva of a weevil or other beetle, by its relatively smaller head and smoother, less wrinkled cuticle; it strikes the observer as a feebler, more helpless creature than a beetle-grub. ”

“ But it is advisable in connection with our present subject to dwell especially on some insects that remain continually under water till they are ready to undergo their final moult and attain the winged state, which they pass entirely in the air. The preparatory instars of such insects are aquatic; the adult instar is aerial. All may-flies, dragon-flies, and caddis-flies, many beetles and two-winged flies, and a few moths thus divide their life-story between the water and the air. ”

“ In any case this region is pushed out during the formation of the pupa within the final larval cuticle, so that the imaginal head with its contained brain, its compound eyes, and its complex feelers, takes its rightful place at the front end of the insect.
The mention of the brain suggests a few brief remarks on the changes in the internal organs during insect transformation. There are no imaginal discs for the nervous system; the brain, nerve-cords and ganglia of the butterfly or bluebottle are the direct outcome of those of the caterpillar or maggot. ”

“ Rocks of Triassic age have yielded beetles and lacewing-flies, while from among Jurassic fossils specimens have been described as representing most of our existing orders, including Lepidoptera, Hymenoptera and Diptera. In Cainozoic rocks fossil insects of nearly six thousand species have been found, which are easily referable to existing families and often to existing genera. We may conclude then, imperfect though our knowledge of extinct insects is, that some of the most complex of insect life-stories were being worked out before the dawn of the Cainozoic era. ”