Studies in the Theory of Descent, Volume I. Weismann August
less completely covered with black than in the male.
IV. Why all Polygoneutic Species are not Seasonally Dimorphic
If we may consider it to be established that seasonal dimorphism is nothing else than the splitting up of a species into two climatic varieties in one and the same locality, the further question at once arises why all polygoneutic species (those which produce more than one annual generation) are not seasonally dimorphic.
To answer this, it will be necessary to go more deeply into the development of seasonal dimorphism. This evidently depends upon a peculiar kind of periodic, alternating heredity, which we might be tempted to identify with Darwin’s “inheritance at corresponding periods of life.” It does not, however, in any way completely agree with this principle, although it presents a great analogy to it and must depend ultimately upon the same cause. The Darwinian “inheritance at corresponding periods of life” – or, as it is termed by Haeckel, “homochronic heredity” – is characterized by the fact that new characters always appear in the individuals at the same stage of life as that in which they appeared in their progenitors. The truth of this principle has been firmly established, instances being known in which both the first appearance of a new (especially pathological) character and its transmission through several generations has been observed. Seasonally dimorphic butterflies also furnish a further valuable proof of this principle, since they show that not only variations which arise suddenly (and which are therefore probably due to purely innate causes) follow this mode of inheritance, but also that characters gradually called forth by the influence of external conditions and accumulating from generation to generation, are only inherited at that period of life in which these conditions were or are effective. In all seasonally dimorphic butterflies which I have been able to examine closely, I found the caterpillars of the summer and winter broods to be perfectly identical. The influences which, by acting on the pupæ, split up the imagines into two climatic forms, were thus without effect on the earlier stages of development. I may specially mention that the caterpillars, as well as the pupæ and eggs of A. Levana, are perfectly alike both in the summer and winter forms; and the same is the case in the corresponding stages of P. Napi and P. Bryoniæ.
I shall not here attempt to enter more deeply into the nature of the phenomena of inheritance. It is sufficient to have confirmed the law that influences which act only on certain stages in the development of the individual, even when the action is cumulative and not sudden, only affect those particular stages without having any effect on the earlier or later stages. This law is obviously of the greatest importance to the comprehension of metamorphosis. Lubbock32 has briefly shown in a very clear manner how the existence of metamorphosis in insects can be explained by the indirect action of varying conditions on the different life-stages of a species. Thus the mandibles of a caterpillar are, by adaptation to another mode of nourishment, exchanged at a later period of life for a suctorial organ. Such adaptation of the various development-stages of a species to the different conditions of life would never give rise to metamorphosis, if the law of homochronic, or periodic, heredity did not cause the characters gradually acquired at a given stage to be transferred to the same stage of the following generation.
The origin of seasonal dimorphism depends upon a very similar law, or rather form, of inheritance, which differs from that above considered only in the fact that, instead of the ontogenetic stages, a whole series of generations is influenced. This form of inheritance may be formulated somewhat as follows: – When dissimilar conditions alternatingly influence a series of generations, a cycle is produced in which the changes are transmitted only to those generations which are acted upon by corresponding conditions, and not to the intermediate ones. Characters which have arisen by the action of a summer climate are inherited by the summer generation only, whilst they remain latent in the winter generation. It is the same as with the mandibles of a caterpillar which are latent in the butterfly, and again make their appearance in the corresponding (larval) stage of the succeeding generation. This is not mere hypothesis, but the legitimate inference from the facts. If it be admitted that my conception of seasonal dimorphism as a double climatic variation is correct, the law of “cyclical heredity,”33 as I may term it – in contradistinction to “homochronic heredity,” which relates only to the ontogenetic stages – immediately follows. All those cases which come under the designation of ‘alternation of generation,’ can obviously be referred to cyclical heredity, as will be explained further on. In the one case the successive generations deport themselves exactly in the same manner as do the successive stages of development of the individual in the other; and we may conclude therefrom (as has long been admitted on other grounds) that a generation is, in fact, nothing else than a stage of development in the life of a species. This appears to me to furnish a beautiful confirmation of the theory of descent.
Now if, returning to questions previously solved, the alternating action of cold in winter and warmth in summer leads to the production of a winter and summer form, according to the law of cyclical heredity, the question still remains: why do we not find seasonal dimorphism in all polygoneutic butterflies?
We might at first suppose that all species are not equally sensitive to the influence of temperature: indeed, the various amounts of difference between the winter and summer forms in different species would certainly show the existence of different degrees of sensitiveness to the modifying action of temperature. But even this does not furnish an explanation, since there are butterflies which produce two perfectly similar34 generations wherever they occur, and which, nevertheless, appear in different climates as climatic varieties. This is the case with Pararga Ægeria (Fig. 23, Plate II.), the southern variety of which, Meione (Fig. 24, Plate II.), is connected with it by an intermediate form from the Ligurian coast. This species possesses, therefore, a decided power of responding to the influence of temperature, and yet no distinction has taken place between the summer and the winter form. We can thus only attribute this different deportment to a different kind of heredity; and we may therefore plainly state, that changes produced by alternation of climate are not always inherited alternatingly, i.e. by the corresponding generations, but sometimes continuously, appearing in every generation, and never remaining latent. The causes which determine why, in a particular case, the one or the other form of inheritance prevails, can be only innate, i.e. they lie in the organism itself, and there is as little to be said upon their precise nature as upon that of any other process of heredity. In a similar manner Darwin admits a kind of double inheritance with respect to characters produced by sexual selection; in one form these characters remain limited to the sex which first acquired them, in the other form they are inherited by both sexes, without it being apparent why, in any particular case, the one or the other form of heredity should take place.
The foregoing explanation may obtain in the case of sexual selection, in which it is not inconceivable that certain characters may not be so easily produced, or even not produced at all, in one sex, owing to its differing from the other in physical constitution. In the class of cases under consideration, however, it is not possible that the inherited characters can be prevented from being acquired by one generation owing to its physical constitution, since this constitution was similar in all the successive generations before the appearance of dimorphism. The constitution in question first became dissimilar in the two generations to the extent of producing a change of specific character, through the action of temperature on the alternating broods of each year, combined with cyclical heredity. If the law of cyclical heredity be a general one, it must hold good for all cases, and characters acquired by the summer generation could never have been also transmitted to the winter generation from the very first.
I will not deny the possibility that if alternating heredity should become subsequently entirely suppressed throughout numerous generations, a period may arrive when the preponderating influence of a long series of summer generations may ultimately take effect upon the winter generation. In such a case the summer characters would appear, instead of remaining latent as formerly. In this manner it may be imagined that at first but few, and later more numerous individuals, approximate to the summer form, until finally the dimorphism entirely disappears, the new form
“On the Origin and Metamorphoses of Insects,” London, 1874.
I at first thought of designating the two forms of cyclical or homochronic heredity as ontogenetic- and phyletic-cyclical heredity. The former would certainly be correct; the latter would be also applicable to alternation of generation (in which actually two or more phyletic stages alternate with each other) but not to all those cases which I attribute to heterogenesis, in which, as with seasonal dimorphism, a series of generations of
When Meyer-Dürr, who is otherwise very accurate, states in his “Verzeichniss der Schmetterlinge der Schweiz,” (1852, p. 207), that the winter and summer generations of