This theory of Sharp's is original, daring, and ingenious, but the loss and re-acquisition of wings which it presupposes is difficult to imagine in large groups during a prolonged evolutionary history, while the sudden appearance of a totally new mode of wing-growth in the offspring of wingless insects would be an extreme example of discontinuity in development.
On the whole the most probable suggestion which can be made as to the origin of 'complete' transformation in insects is that the instar in which wings were first visible externally became later and later in the course of the evolution of the more highly organised groups. In this way a gradual transition from the exopterygote to the endopterygote type of life-story is at least conceivable. It will be remembered that a may-fly ([p. 33]) undergoes a moult after acquiring functional wings, emerging into the air as a 'sub-imago.' In not a few endopterygote insects, the pupa shows more or less activity, swimming through water intermittently (gnats) or just before the imago has to emerge (caddis-flies); working its way out of the ground (crane-flies) or coming half-way out of its cocoon (many moths). The pupa of the higher insects almost certainly corresponds with the may-fly's sub-imago, and the facts just recalled as to remnants of pupal activity suggest that in the ancestors of endopterygote insects what is now the pupal instar was represented by an active nymphal or sub-imaginal stage, possibly indeed by more than one stage, as Packard and other writers have stated that pupae of bees and wasps undergo two or three moults before the final exposure of the imago. Such an early pupal instar has been defined as a 'pro-nymph' or a 'semi-pupa.' Examples have been given of the exceptional passive condition of the penultimate instar in Exopterygota. The instars preceding this presumably had originally outward wing-rudiments in all insect life-histories, and the endopterygote condition was attained by the postponement of the outward appearance of these to successively later stages. The leg and wing rudiments of the male coccid ([pp. 20-1]) beneath the cuticle of the second instar are strictly comparable to imaginal buds, and these are present in one instar of what is generally regarded as an exopterygote life-history. The first instar in all insects has no visible wing-rudiments, but when they grow outwardly from the body, they necessarily become covered with cuticle, so that they must be visible after the first moult. There is no supreme difficulty in supposing that the important change was for these early rudiments to become sunk into the body, so that the cuticle of the second, and, later, of the third and succeeding instars, showed no outward sign of their presence. This suggestion is confirmed by Heymons' ([1896], [1907]) observation of the occasional appearance of outward wing-rudiments on the thoracic segments of a mealworm, the larva of the beetle Tenebrio molitor, and by F. Silvestri's discovery [(1905)] of a 'pro-nymph' stage with short external wing-rudiments between the second larval and the pupal instars of the small ground-beetle Lebia scapularis. Whatever may be the exact explanation of these abnormalities, they show that in the life-story of the higher insects outward wing-rudiments may even yet appear before the pupal stage, confirming our belief that such appearance is an ancestral character. The inward growth of these wing-rudiments may well have been correlated with a difference in form between the newly-hatched insect and its parent. As this difference persisted until a constantly later stage, and the pre-imaginal instar became necessarily a stage for reconstruction, the present condition of complete metamorphosis in the more highly organised orders was finally attained.
To explain satisfactorily these complex life-stories is however admittedly a difficult task. The acquisition of wings is, as we have seen, a dominating feature in them all, but if we try to go yet a step farther back and speculate on the origin of wings in the most primitive exopterygote insects, the task becomes still more difficult. Many years ago [Gegenbaur (1878)] was struck by the correspondence of insect wings to the tracheal gills of may-fly larvae, which are carried on the abdominal segments somewhat as wings are on the thoracic segments. But Börner has recently [(1909)] brought forward evidence that these abdominal gills really correspond serially with legs. Moreover Gegenbaur's theory suggests that the ancestral insects were aquatic, whereas the presence of tubes for breathing atmospheric air in well-nigh all members of the class, and the fact that aquatic adaptations, respiratory and otherwise, in insect-larvae are secondary force the student to regard the ancestral insects as terrestrial. It is indeed highly probable that insects had a common origin with aquatic Crustacea, but all the evidence points to the ancestors of insects having become breathers of atmospheric air before they acquired wings. How the wings arose, what function their precursors performed before they became capable of supporting flight, we can hardly even guess.
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. 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. But the light which research has thrown on the nature of these wonderful life-stories, the demonstration of the unseen presence and growth within the insect, during its time of preparation among strange surroundings, of the organs required for service in the coming life amid its native air, confirm surely the intuition of the old-time students, who saw in these changes, so familiar and yet so wonderful, a parable and a prophecy of the higher nature of man.
OUTLINE CLASSIFICATION OF INSECTS
Class INSECTA or HEXAPODA.
Sub-class A, Apterygota.
- Order
- Thysanura (Bristle-tails).
- Collembola (Spring-tails).