Further facts can be adduced in support of the view that in these processes it is the constitution of the species, and not the external agency (warmth), which plays the chief part. The latter, as Darwin has strikingly expressed it, rather performs the function of the spark which ignites a combustible substance, whilst the character of the combustion depends upon the nature of the explosive material. Were this not the case, increased warmth would always change a given colour[29] in the same manner in all butterflies, and would therefore always give rise to the production of the same colour. But this does not occur; Polyommatus Phlæas, for example, becoming black in the south, whilst the red-brown Vanessa Urticæ becomes black in high northern latitudes, and many other cases well known to entomologists might be adduced.[30] It indeed appears that species of similar physical constitution, i.e., nearly allied species, under similar climatic influences, change in an analogous manner. A beautiful example of this is furnished by our Pierinæ. Most of the species display seasonal dimorphism; as, for instance, Pieris Brassicæ, Rapæ, Napi, Krueperi, and Daplidice, Euchloe Belia and Belemia, and Leucophasia Sinapis, in all of which the difference between the winter and the summer forms is of a precisely similar nature. The former are characterized by a strong black dusting of the base of the wings, and by a blackish or green sprinkling of scales on the underside of the hind wings, while the latter have intensely black tips to the wings, and frequently also spots on the fore-wings.
Nothing can prove more strikingly, however, that in such cases everything depends upon the physical constitution, than the fact that in the same species the males become changed in a different manner to the females. The parent-form of Pieris Napi (var. Bryoniæ) offers an example. In all the Pierinæ secondary sexual differences are found, the males being differently marked to the females; the species are thus sexually dimorphic. Now the male of the Alpine and Polar var. Bryoniæ, which I conceive to be the ancestral form, is scarcely to be distinguished, as has already been mentioned, from the male of our German winter form (P. Napi, var. Vernalis), whilst the female differs considerably.[31] The gradual climatic change which transformed the parent-form Bryoniæ into Napi has therefore exerted a much greater effect on the female than on the male. The external action on the two sexes was exactly the same, but the response of the organism was different, and the cause of the difference can only be sought for in the fine differences of physical constitution which distinguish the male from the female. If we are unable to define these differences precisely, we may nevertheless safely conclude from such observations that they exist.
I have given special prominence to this subject because, in my idea, Darwin ascribes too much power to sexual selection when he attributes the formation of secondary sexual characters to the sole action of this agency. The case of Bryoniæ teaches us that such characters may arise from purely innate causes; and until experiments have decided how far the influence of sexual selection extends, we are justified in believing that the sexual dimorphism of butterflies is due in great part to the differences of physical constitution between the sexes. It is quite different with such sexual characters as the stridulating organs of male Orthoptera which are of undoubted importance to that sex. These can certainly be attributed with great probability to sexual selection.
It may perhaps not be superfluous to adduce one more similar case, in which, however, the male and not the female is the most affected by climate. In our latitudes, as also in the extreme north, Polyommatus Phlæas, already so often mentioned, is perfectly similar in both sexes in colour and marking; and the same holds good for the winter generation of the south. The summer generation of the latter, however, exhibits a slight sexual dimorphism, the red of the fore wings of the female being 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. Lubbock[32] 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.