In Smerinthus Tiliæ, Ocellatus and Populi also, the greatest larval variability is shown only in the last stage, the preceding stages being very constant. These cases by no means depend upon the marking of the young stages being simpler and therefore being less capable of varying. The reverse case also occurs. In a somewhat similar manner as the young of the tapir and wild hog are striped, while the adult animals are plainly coloured, the young caterpillars of Saturnia Yamamai possess longitudinal black lines on a yellow ground, while as early as in the second stage a simple green colour appears in the place of this complicated but perfectly constant marking. If the young stages are so frequently constant, this rather depends upon the fact that the transference of a new character to these stages not only takes place gradually, but also with continually diminishing energy, in a manner somewhat similar to physical motion, which continually diminishes in speed by the action of resistance till it is completely arrested. This constancy of the younger stages may further be due to the circumstance that the characters would only be transferred when they had become fixed in the last stage, and were consequently no longer variable. The transferred characters may thus have acquired a greater regularity, i.e. a less degree of variability, than they possessed at their first origination. Extensive investigations in this special direction must be made if the precise laws, in accordance with which the backward transference of new characters takes place, are to be discovered. By such researches only should we arrive with certainty at the causes which determine the lesser variability of the young larval stages.
It may also occur that the early stages are variable, whilst the later stages are constant, although this case appears to happen less frequently. Thus, the caterpillars of Gastropacha Quercifolia vary considerably in the second stage but are constant at a later period, and the same is the case with Spilosoma Urticæ, which in the second stage may be almost considered to be dimorphic, but which subsequently becomes constant.
Cases in which the first stage is variable appear to be of the least frequent occurrence. I know of only one such instance, viz., Anceryx Pinastri, of which the newly hatched larvæ ([Pl. VI]., Fig. 53) show considerable differences in the brownish-black crescentic spots. The second ([Fig. 54]), third, and fourth stages are then tolerably constant, while the fifth stage again is very variable.
An instance of this kind can be easily explained by two waves of variation, the first of which now affects only the first stage, while the second has just commenced to affect the fifth stage. Such a supposition is not opposed to any theoretical considerations, but rather has much probability in its favour, since we know that species are from time to time subject to be remodelled; and further, that the coalescence of several stages of phyletic development in the ontogeny of one and the same species (see p. [226], development of the genus Deilephila) shows that during the backward transference of one character, new characters may appear in the last stage of the ontogeny, and indeed very frequently at a time when the next youngest character has not been transferred back so far as to the first stage.
That this secondary variability is to a certain extent brought about by the conflict between the old and new characters, the latter striving to suppress the former, is shown by the caterpillar of Saturnia Carpini which I have observed for many years from this point of view, and than which I do not know a more beautiful illustration.
When these larvæ leave the egg they are black, but in the adult state are almost bright green—this at least being the case in a local form which, from the district in the vicinity of Genoa where it is found, I will designate as the var. Ligurica. Now whilst these two extreme stages of development are relatively constant, the intermediate stages show a variability which becomes greater the nearer the last stage is approached, this variation in the marking depending simply on the struggle between the green colour and the more anciently inherited black. In this manner there arises, especially in the fourth stage of the German local form, an incredible mixture of the most diverse markings, all of which can, however, be very easily explained from the foregoing point of view.
The simpler and, as I am inclined to believe, the older form of the transformation is presented to us in the local variety Ligurica. In the last stage, when 7.5 centimeters long, this form is of a beautiful bright green colour without any trace of black marking[174] ([Pl. VIII]., Fig. 77). The colour of the six orange warts which are situated on each segment is also similar in all specimens, so that this stage is perfectly constant.
Our German S. Carpini shows different characters in the fifth stage. It is true that individual specimens occur which are entirely green without any black, but these are rare; the majority possess a more or less broad black ring encircling the middle of each segment ([Pl. VIII]., Figs. 78 and 79). Those specimens in which the black ring has become broken up into large or small spots surrounding the base of the warts constitute intermediate forms ([Fig. 80]). The last stage of the German local form, unlike that of the Genoese local form, is therefore very variable.
The two forms, moreover, do not simply differ in being more or less advanced in phyletic development, but also in several other points. As it is of great theoretical interest to show that a species can develop local differences only in the stage of larva, I will here subjoin the plain facts.
The differences consist in that the Genoese local form goes through five moults whilst the German local form, like most caterpillars, has only four moults. Further, in the Genoese form the light green, which is also possessed by the German form in the fourth stage, when it once appears, is retained to the end of the larval development, whilst in the fifth stage of the German form this colour is replaced by a dull greyish-green (compare [Figs. 77 and 78]). There is further a very considerable difference in the earlier stages which shows that the phyletic transforming process has taken a quite independent course in the two forms. Since the struggle between the green and black—retaining this idea—appears to be quite finished in the last stage of the Genoese form, we should expect that the new colour, green, would now also have encroached further upon the younger stages than in the German form. Nevertheless, this is not the case, but quite the reverse happens, the black maintaining its ground longer in the Italian than in the German form.