Let us take the species of one genus, e.g. those of Deilephila. The imagines of our European species—as far as we know—all live in precisely the same manner; they all fly at twilight,[173] showing a preference for the same flowers and very often frequenting the same spots, so that in the haunts of one species the others are almost always to be met with, supposing them to occur in the same locality. They conceal themselves by day in similar places, and are attacked by similar foes.
It is quite different with the caterpillars. These, even in the case of the most closely allied species, live under different conditions, as appears from the fact that they feed on different plants. The latter can, however, produce changes both directly and indirectly. The larvæ may acquire adaptive colours and markings, and these would vary in accordance with the colour and structure of the food-plant; or they may become brightly coloured as a sign of distastefulness in cases where they are inedible. Then again the colour of the soil on which the larvæ live would act upon their colours making these adaptive. Certain habits of the caterpillars may also be dependent upon the nature of their food-plants. Thus, e.g. Deilephila Hippophaës feeds only at night, and conceals itself by day under moss and among the leaves at the base of the food-plant; but D. Euphorbiæ could not acquire such a habit, because Euphorbia Cyparissias generally grows on arid soil which is poor in vegetation, and which therefore affords no concealment, and furthermore, because a caterpillar, as long as it continues to feed, cannot, and as a matter of fact does not, ever wander far from its food-plant. A habit of concealment by burying in the earth also, such for example as occurs in Acherontia Atropos, could not be acquired by D. Euphorbiæ, because its food-plant generally grows on hard, dry, and stony ground.
In addition to these considerations, the foes would be different according as the caterpillar lived on plants which formed dense thickets covering large extents of the shore (Hippophae) or grew isolated on dry hillocks and declivities where the herbage was scanty or altogether absent; or again, according as the insect, in conjunction with such local differences, fed by day or had acquired the habit of feeding only by night. It must in fact be admitted that new and improved adaptations, or, in more general terms, that inducements to change, when depending on the environment, must be more frequently dissimilar for larvæ than for the imagines. We must accordingly expect to find actual change, or that condition of variability which may be regarded as initiative to change, occurring more commonly in larvæ than in perfect insects.
Since facts are in complete accordance with the results of these à priori considerations we may also venture to conclude that the basis of the considerations is likewise correct, viz., the supposition that the changes of colour and marking in caterpillars, pupæ, and imagines result from external influences only.
This must not be taken as signifying that the single stages of the larval development are also only able to change through the action of external influences. The larval stages are correlated with each other, as has already been shown (see the previous essay): new characters arise in the adult caterpillar at the last stage and are then gradually transferred back to the younger stages quite independently of external influences, this recession being entirely brought about by the laws of correlation. Natural selection here only exerts a secondary action, since it can accelerate or retard this transference, according as the new characters are advantageous or disadvantageous to the younger stages.
Now as considerable individual differences appear in the first acquisition of a new character with respect to the rapidity and completeness with which the individuals acquire such a character, the same must obtain for the transference of an improvement acquired in the last stage to the next younger stage. The new character would be acquired by different individuals in different degrees and at different rates—it would have, to a certain extent, to struggle with the older characters of the stage; in brief, the younger stage would become variable.
Variability of this kind might well be designated as secondary, in contradistinction to primary variability; the latter (primary) depends upon an unequal reaction of the individual organisms to external influences, the former (secondary) results from the unequal strength and rate of the action of the innate laws of growth governing the organism. In both cases alike exceeding variability may occur, but the causes producing this variability are dissimilar.
The different stages of larval development would thus frequently display independent variability in a manner similar to the pupal or imaginal stages, since they can show individual variability while the other stages of development remain constant. This appearance of independent variability in the different stages of the larval development, however, is in truth deceptive—we have here in fact a kind of wave of variability, which passes downwards through the developmental stages, becoming gradually weaker, and finally dying out completely.
In accordance with this, we very frequently find that only the last or two last stages are variable, while the younger stages are constant. Thus in Macroglossa Stellatarum, the larvæ are constant in the first, second, and third stages, but become variable in the fourth, and in the fifth stage first show that high degree of variability which has already been described in detail (See. [Pl. III]., Figs. 3–12). The larvæ, of Vanessa Cardui also, according to my notes, are extremely constant in the first four stages in spite of their complicated marking, but become variable in the fifth stage, although to no very great extent.