If we turn to the groups of families we find a considerably increased incongruence; complete agreement is here again rather the exception, and it further happens in these cases that it is always the larvæ which, to a certain extent, remain at a lower grade, and which form well defined families; but these can seldom be associated into groups of a higher order having a common character, as in the case of the imagines (Rhopalocera).
After having thus collected (so far as I am able) the facts, we have now to attempt their interpretation, and from the observed congruence and incongruence of form-relationship of the two stages to endeavour to draw a conclusion as to the underlying causes of the transformations.
It is clear at starting that all cases of incongruence can only be the expression or the consequence of a phyletic development which has not been exactly parallel in the two stages of larva and imago—that one stage must have changed either more rapidly or more slowly than the other. An “unequal phyletic development” is thus the immediate cause of incongruence.
Thus, the occurrence of different larvæ in species of which the imagines have remained alike may be simply understood as cases in which the imago only has experienced a change—has taken a forward step in phyletic development, whilst the larvæ have remained behind. If we conceive this one-sided development to be repeated several times, there would arise two larval forms as widely different as those of Deilephila Nicæa, and Euphorbiæ, whilst the imagines, as is actually the case in these species, would remain the same.
The more commonly occurring case in which one stage has a greater form-divergence than the other, is explicable by the one stage having changed more frequently or more strongly than the other.
The explanation of the phenomena thus far lies on the surface, and it is scarcely possible to advance any other; but why should one stage become changed more frequently or to a greater extent than the other? why should one portion be induced to change more frequently or more strongly than another? whence come these inducements to change? These questions bring us to the main point of inquiry:—Are the causes which give rise to these changes internal or external? Are the latter the result of a phyletic vital force, or are they only due to the action of the external conditions of life?
Although an answer to this question will be found in the preceding essay, I will not support myself on the results there obtained, but will endeavour to give another solution of the problem on fresh grounds. The answer will indeed be the same as before:—A phyletic force must be discountenanced, since in the first place it does not explain the phenomena, and in the second place the phenomena can be well explained without its assumption.
The admission of a phyletic vital force does not explain the phenomena. The assumption that there is a transforming power innate in the organism indeed agrees quite well with the phenomenon of congruence, but not with that of incongruence. Since a large number of cases of the latter depend upon the fact that the larvæ are more frequently influenced by causes of change than their imagines, or vice versâ, how can this be reconciled with such an internal force? On this assumption would not each stage of a species be compelled to change, if not contemporaneously at least successively, with the same frequency and intensity, by the action of an innate force? and how by means of the latter can there ever result a greater form-divergence in the larvæ than in the imagines?
It is delusive to believe that these unequal deviations can be explained by assuming that the phyletic force acts periodically. Granting that it does so, and that the internal power successively compels the imago, pupa, and finally the larva to change, there would then pass a kind of wave of transformation over the different stages of the species, as was actually shown above to be the case in the single larval stages. The only possible way of explaining the unequal distances between larvæ and imagines would therefore be to assume that two allied groups, e.g. species, were not contemporaneously affected by the wave, so that at a certain period of time the imago alone of one species had become changed, whilst in the other species the wave of transformation had also reached the larva. In this case the imagines of the two species would thus appear to be more nearly related than their larvæ.