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 thus gaining ascendency and the species becoming once more monomorphic. Such a supposition is indeed capable of being supported by some facts, an observation on A. Levana apparently contradicting the theory having been already interpreted in this sense. I refer to the fact that whilst some butterflies of the winter generation emerge in October as Prorsa, others hibernate, and appear the following spring in the Levana form. The winter form of Pieris Napi also no longer preserves, in the female sex, the striking coloration of the ancestral form Bryoniæ, a fact which may indicate the influencing of the winter generation by numerous summer generations. The double form of the spring generation of Papilio Ajax can be similarly explained by the gradual change of alternating into continuous heredity, as has already been mentioned. All these cases, however, are perhaps capable of another interpretation; at any rate, the correctness of this supposition can only be decided by further facts.
Meanwhile, even if we suppose the above explanation to be correct, it will not apply to the absence of seasonal dimorphism in cases like that of Pararga Ægeria and Meione, in which only one summer generation appears, so that a preponderating inheritance of summer characters cannot be admitted. Another explanation must thus be sought, and I believe that I have found it in the circumstance that the butterflies named do not hibernate as pupæ but as caterpillars, so that the cold of winter does not directly influence those processes of development by which the perfect insect is formed in the chrysalis. It is precisely on this point that the origin of those differences of colour which we designate as the seasonal dimorphism of butterflies appears to depend. Previous experiments give great probability to this statement. From these we know that the eggs, caterpillars, and pupæ of all the seasonally dimorphic species experimented with are perfectly similar in the summer and winter generations, the imago stage only showing any difference. We know further from these experiments, that temperature-influences which affect the caterpillars never entail a change in the butterflies; and finally, that the artificial production of the reversion of the summer to the winter form can only be brought about by operating on the pupæ.
Since many monogoneutic species now hibernate in the caterpillar stage (e.g. Satyrus Proserpina, and Hermione, Epinephele Eudora, Furtina, Ithonus, Hyperanthus, Ida, &c.), we may admit that during the glacial period such species did not pass the winter as pupæ. As the climate grew warmer, and in consequence thereof a second generation became gradually interpolated in many of these monogoneutic species, there would ensue (though by no means necessarily) a disturbance of the winter generation, of such a kind that the pupæ, instead of the caterpillars as formerly, would then hibernate. It may, indeed, be easily proved à priori that whenever a disturbance of the winter generation takes place it only does so retrogressively, that is to say – species which at one time pass the winter as caterpillars subsequently hibernate in the egg, while those which formerly hibernate as pupæ afterwards do so as caterpillars. The interpolation of a summer generation must necessarily delay till further towards the end of summer, the brood about to hibernate; the remainder of the summer, which serves for the development of the eggs and young caterpillars, may possibly under these conditions be insufficient for pupation, and the species which hibernated in the pupal state when it was monogoneutic, may perhaps pass the winter in the larval condition after the introduction of the second brood. A disturbance of this kind is conceivable; but it is certain that many species suffer no further alteration in their development than that of becoming digoneutic from monogoneutic. This follows from the fact that hibernation takes place in the caterpillar stage in many species of the sub-family Satyridæ which are now digoneutic, as well as in the remaining monogoneutic species of the same sub-family. But we cannot expect seasonal dimorphism to appear in all digoneutic butterflies the winter generation of which hibernates in the caterpillar form, since the pupal stage in these species experiences nearly the same influences of temperature in both generations. We are hence led to the conclusion that seasonal dimorphism must arise in butterflies whenever the pupæ of the alternating annual generations are exposed throughout long periods of time to widely different regularly recurring changes of temperature.
The facts agree with this conclusion, inasmuch as most butterflies which exhibit seasonal dimorphism hibernate in the pupa stage. Thus, this is the case with all the Pierinæ, with Papilio Machaon, P. Podalirius, and P. Ajax, as well as with Araschnia Levana. Nevertheless, it cannot be denied that seasonal dimorphism occurs also in some species which do not hibernate as pupæ but as caterpillars; as, for instance, in the strongly dimorphic Plebeius Amyntas. But such cases can be explained in a different manner.
Again, the formation of a climatic variety – and as such must we regard seasonally dimorphic forms – by no means entirely depends on the magnitude of the difference between the temperature which acts on the pupæ of the primary and that which acts on those of the secondary form; it rather depends on the absolute temperature which the pupæ experience. This follows without doubt from the fact that many species, such as our common Swallow-tail (Papilio Machaon), and also P. Podalirius, in Germany and the rest of temperate Europe, show no perceptible difference of colour between the first generation, the pupæ of which hibernate, and the second generation, the pupal period of which falls in July, whereas the same butterflies in South Spain and Italy are to a small extent seasonally dimorphic. Those butterflies which are developed under the influence of a Sicilian summer heat likewise show climatic variation to a small extent. The following consideration throws further light on these conditions. The mean summer and winter temperatures in Germany differ by about 14.9° R.; this difference being therefore much more pronounced than that between the German and Sicilian summer, which is only about 3.6° R. Nevertheless, the winter and summer generations of P. Podalirius are alike in Germany, whilst the Sicilian summer generation has become a climatic variety. The cause of this change must therefore lie in the small difference between the mean summer temperatures of 15.0° R. (Berlin) and 19.4° R. (Palermo). According to this, a given absolute temperature appears to give a tendency to variation in a certain direction, the necessary temperature being different for different species. The latter statement is supported by the facts that, in the first place, in different species there are very different degrees of difference between the summer and winter forms; and secondly, many digoneutic species are still monomorphic in Germany, first becoming seasonally dimorphic in Southern Europe. This is the case with P. Machaon and P. Podalirius, as already mentioned, and likewise with Polyommatus Phlæas. Zeller in 1846–47, during his journey in Italy, recognized as seasonally dimorphic in a small degree a large number of diurnal Lepidoptera which are not so in our climate.35
In a similar manner the appearance of seasonal dimorphism in species which, like Plebeius Amyntas, do not hibernate as pupæ, but as caterpillars, can be simply explained by supposing that the winter generation was the primary form, and that the increase in the summer temperature since the glacial period was sufficient to cause this particular species to become changed by the gradual interpolation of a second generation. The dimorphism of P. Amyntas can, nevertheless, be explained in another manner. Thus, there may have been a disturbance of the period of development in the manner already indicated, the species which formerly hibernated in the pupal stage becoming subsequently disturbed in its course of development by the interpolation of a summer generation, and hibernating in consequence in the caterpillar state. Under these circumstances we must regard the present winter form (var. Polysperchon) as having been established under the influence of a winter climate, this form, since the supposed disturbance in its development, having had no reason to become changed, the spring temperature under which its pupation now takes place not being sufficiently high. The interpolated second generation on the other hand, the pupal period of which falls in the height of summer, may easily have become formed into a summer variety.
This latter explanation agrees precisely with the former, both starting with the assumption that in the present case, as in that of A. Levana and the Pierinæ, the winter form is the primary one, so that the dimorphism proceeds from the said winter form and does not originate the winter but the summer form, as will be explained. Whether the winter form has been produced by the action of the winter or spring temperature is immaterial in judging single cases, inasmuch as we are not in a position to state what temperature is necessary to cause any particular species to become transformed.
The reverse case is also theoretically conceivable, viz., that in certain species the summer form was the primary one, and by spreading northwards a climate was reached which still permitted the production of two generations, the pupal stage of one generation being exposed to the cold of winter, and thus giving rise to the production of a secondary winter form. In such a case hibernation in the pupal state would certainly give rise to seasonal dimorphism. Whether these conditions actually occur, appears to me extremely doubtful; but it may at least be confidently asserted that the first case is of far more frequent occurrence. The beautiful researches of Ernst Hoffmann36 furnish strong evidence for believing that the great majority of the European butterflies have immigrated, not from the south, but from Siberia. Of 281 species, 173 have, according to Hoffmann, come from Siberia, 39 from southern Asia, and only 8 from Africa, whilst during the greatest cold of the glacial period, but very few or possibly no species existed north of the Alps. Most of the butterflies now found in Europe have thus, since their immigration, experienced a gradually increasing warmth. Since seasonal dimorphism has been developed in some of these species, the summer form must in all cases have been the secondary one, as the experiments upon the reversion of Pieris Napi and Araschnia Levana have also shown.
All the seasonally dimorphic butterflies known to me are found in Hoffmann’s list of Siberian immigrants, with the exception of two species, viz., Euchloe Belemia, which is cited as an African immigrant, and Pieris Krueperi, which may have come through Asia Minor, since at the present time it has not advanced farther west than Greece. No considerable change of climate can be experienced by migrating from east to west, so that the seasonal dimorphism of Pieris Krueperi can only depend on a cause similar to that which affected the Siberian immigrants, that is, the gradual increase of temperature in the northern hemisphere since the glacial period. In this species also, the winter form must be the primary one. In the case of E. Belemia, on the other hand, the migration northwards from Africa certainly indicates removal to a cooler climate, which may have originated a secondary winter form, even if nothing more certain can be stated. We know nothing of the period of migration into southern Europe; and even migration without climatic change is conceivable, if it kept pace with the gradual increase of warmth in the northern hemisphere since the glacial epoch. Experiments only would in this case be decisive. If the summer generation, var. Glauce, were the primary form, it would not be possible by the action of cold on the pupæ of this brood to produce the winter variety Belemia, whilst, on the other hand, the pupæ of the winter generation by the influence of warmth would be made to revert more or less completely to the form Glauce. It is by no means to be understood that the species would actually comport itself in this manner. On the contrary, I am of opinion that in this case also, the winter form is primary. The northward migration (from Africa to south Spain) would be quite insufficient, and the winter form is now found in Africa as well as in Spain.