Studies in the Theory of Descent, Volume I

In accordance with the idea first introduced into science by Rudolph Leuckart, we have hitherto understood heterogenesis to be only the alternation of dissimilar sexual generations. From this point of view the reproduction of Leptodora can be as little ascribed to heterogenesis as can that of Aphis or Daphnia, although the apparent agamic reproduction of the winter and a portion of the summer generation is undoubtedly parthenogenesis and not propagation by nursing.48 As has already been said, however, I would attribute no fundamental importance to the criterion of agamic reproduction – the more especially because we are ignorant of the physiological significance of the two modes of propagation; and further, because this principle of classification is entirely external, and only valuable in so far as no better one can be substituted for it. A separation of the modes of cyclical propagation according to their genesis appears to me – especially if practicable – not alone to be of greater value, but the only correct one, and for this the knowledge of the origin of seasonal dimorphism seems to me to furnish a possible method.

If, as was indicated above, we designate as metagenesis (in the narrow sense) all those cases in which it must be admitted that a series of differently aged phyletic stages have furnished the points of departure, and as heterogenesis those cases in which similar phyletic stages have been compelled to produce a cycle of generations by the periodic action of external influences, it is clear that the scope of heterogenesis is by this means considerably extended, and at the same time sharply and precisely defined.

Under heterogenesis then is comprised, not only as heretofore the reproduction of Ascaris nigrovenosa, of Leptodora appendiculata, and of the cattle-lice, but also that of the Aphides, Coccidæ, Daphniidæ, Rotatoria, and Phyllopoda, and, in short, all those cases in which we can determine the former identity of the two kinds of generations from their form, anatomical structure, and mode of reproduction. This conclusion is essentially supported by a comparison of the most closely allied species. Thus, for instance, when we see the genus Aphis and its allies related on all sides to insects which propagate sexually in all generations, and when we further observe the great similarity of the whole external and internal structure in the two kinds of generations of Aphis, we are forced to the conjecture that the apparent asexual reproduction of the Aphidæ is in reality parthenogenesis, i.e., that it has been developed from sexual reproduction. Neither can it be any longer disputed that in this case, as well as in that of Leptodora and other Daphniidæ, the same female alternately propagates parthenogenetically, and produces eggs requiring fertilization. This was established by Von Heyden49 some years ago, in the case of Lachnus Querci, and has been since confirmed by Balbiani.50

There can be no doubt that in all these cases the cycle of generations has been developed from phyletically similar generations. But instances are certainly conceivable which present themselves with less clearness and simplicity. In the first place, we do not know whether parthenogenesis may not finally settle down into complete asexual reproduction. Should this be the case, it might be possible that from heterogenesis a mode of propagation would ultimately arise, which was apparently indistinguishable from pure metagenesis. Such a state of affairs might result, if the generations settling into asexual reproduction (as, for instance, the plant-lice), at the same time by adaptation to varying conditions of life, underwent considerable change of structure, and entered upon a metamorphosis to some extent retrogressive. We should then be inclined to regard these generations as an earlier phyletic stage, whilst, in fact, they would be a later one, and the idea of metagenesis would thus have been formed after the manner of heterogenesis.

On the other hand, it is equally conceivable that heterogenesis may have been developed from true metagenesis in the case of larvæ which, having acquired the faculty of asexual propagation, are similar in function to sexually mature insects. This possibility is not at first sight apparent. If the nursing-larvæ of the Cecidomyiæ were as much like the sexual insects as are the young Orthoptera to the sexually mature forms, we should not know whether to regard them as degraded sexual insects, or as true larvæ which had attained the power of asexual propagation. Their propagation would be considered to be parthenogenesis; and as it could not be denied that heterogenesis was here manifest, the mode of development of their particular kind of propagation might be proved, i.e., it might be demonstrated, that the generations now parthenogenetic were formerly mere reproductive larval stages.

I have only offered these last observations in order to show on what uncertain ground we are still standing with regard to this subject whenever we deal with the meaning of any particular case, and how much still remains to be done. It appears certain that the two forms of cyclical propagation, heterogenesis and metagenesis, originate in entirely distinct ways, so that it must be admitted that, under these circumstances, the idea of the existing conditions respecting the true genesis may possibly be erroneous. To indicate the manner in which the cyclical mode of propagation has arisen in any single case, would only be possible by a searching proof and complete knowledge of existing facts in addition to experiments.

VI. General Conclusions

I shall not here give a repetition and summary of the results arrived at with respect to seasonal dimorphism, but rather the general conclusions derived from these results; and, at the same time, I may take the opportunity of raising certain questions which have not hitherto found expression, or have been but briefly and casually stated.

It must, in the first place, be admitted that differences of specific value can originate through the direct action of external conditions of life only. Of the truth of this proposition there can be no doubt, after what has been above stated concerning the difference between the two forms of any seasonally dimorphic species. The best proof is furnished by the older systematists, to whom the genetic relationship of the two forms was unknown, and who, with unprejudiced taxonomy, in many cases indicated their distinctness by separate specific names. This was the case with Araschnia Levana and Prorsa, Euchloe Belia and Ausonia, E. Belemia and Glauce, Plebeius Polysperchon and Amyntas. In the presence of these facts it can scarcely be doubted that new species can be formed in the manner indicated; and I believe that this was and is still the case, with butterflies at least, to a considerable extent; the more so with these insects, because the striking colours and markings of the wings and body, being in most cases without biological significance, are useless for the preservation of the individual or the species, and cannot, therefore, be objects of natural selection.

Darwin must have obtained a clear insight into this, when he attempted to attribute the markings of butterflies to sexual and not to natural selection. According to this view, every new colour or marking first appears in one sex accidentally,51 and is there fixed by being preferred by the other sex to the older coloration. When the new ornamentation becomes constant (in the male for example), Darwin supposes that it becomes transferred to the female by inheritance, either partially or completely, or not at all; so that the species, therefore, remains more or less sexually dimorphic, or (by complete transference) becomes again sexually monomorphic.

The admissibility of such different, and, to a certain extent, arbitrarily limited inheritance, has already been acknowledged. The question here concerned is, whether Darwin is correct when he in this manner attributes the entire coloration of butterflies to sexual selection. The origin of seasonal dimorphism appears to me to be against this view, howsoever seductive and grand the latter may seem. If differences as important as those which exist between the summer and winter forms of many butterflies can be called forth by the direct action of a changed climate, it would be extremely hazardous to attribute great importance to sexual selection in this particular case.

The principle of sexual selection appears to me to be incontestible, and I will not deny that it is also effective in the case of butterflies; but I believe that as a final explanation of colour this agency can be dispensed with, inasmuch as we see that considerable changes of colour can occur without the influence of sexual selection.52

The question now arises, how far does the transforming influence of climate extend? When a species has become transformed by climatic change to such an extent that its new form possesses the systematic value of a new species, does it return to its older form by removal to the old climatic conditions? or would it under these circumstances become again transformed in a new manner? This question is not without importance, inasmuch as in the first case climatic influences would be of little value in the formation of species, and there would result at most only a fluctuation between two extremes. In the same manner as in seasonally dimorphic species the summer and winter forms now alternate with each other every year, so would the forms produced by warmth and cold then alternate in the greater periods of the earth’s history. Other groups of animals are certainly changed by the action of different climatic influences; but in butterflies, as I believe I have proved, temperature plays the chief part, and as this only oscillates between rather narrow limits, it admits of no great differences of coloration.

The question thus suggests itself, whether species of butterflies only oscillate between two forms, or whether climatic change, when sufficiently great to produce variation, does not again originate a new form. Inasmuch as the reversion experiments with seasonally dimorphic butterflies appear to correspond with the latter view, I believe that this must be admitted. I am of opinion that an old form never again arises through change of climate, but always a new one; so that a periodically recurring change of climate is alone sufficient, in the course of a long period of time, to admit of new species arising from one another. This, at least, may be the case with butterflies.

My views rest essentially upon theoretical considerations. It has already been insisted upon, as results immediately from the experiments, that temperature does not act on the physical constitution of the individual in the same manner as acid or alkali upon litmus paper, i.e., that one and the same individual does not produce this or that coloration and marking according as it is exposed to warmth or cold; but rather that climate, when it influences in a similar manner many succeeding generations, gradually produces such a change in the physical constitution of the species that this manifests itself by other colours and markings. Now when this newly acquired physical constitution, established, as we may admit, throughout a long series of generations, is again submitted to a constant change of climate, this influence, even if precisely similar to that which obtained during the period of the first form of the species, cannot possibly reproduce this first form. The nature of the external conditions may be the same, but not so the physical constitution of the species. Just in the same manner as a Pieris (as has been already shown), a Lycæna, or a Satyrus, produces quite different varieties under the transforming influence of the same climate, so must the variation originating from the transformed species of our present case after the beginning of the primary climate be different from that primary form of the species, although perhaps in a less degree. In other words, if only two different climates alternated with each other during the earth’s geological periods, every species of butterfly submitted to these changes of climate would give rise to an endless series of different specific forms. The difference of climate would in reality be greater than supposed, and for any given species the climatic variation would not only occur through the periodic shifting of the ecliptic, but also through geological changes and the migrations of the species itself, so that a continuous change of species must have gone on from this sole cause of alternation of climate. When we consider that many species elsewhere extinct have become locally preserved, and when, further, to these we add those local forms which have arisen by the prevention of crossing (amixia), and finally take into consideration the important effects of sexual selection, we can no longer be astonished at the vast numbers of species of butterflies which we now meet with on the earth.

Should any one be inclined to conclude, from my reversion experiments with seasonally dimorphic butterflies, that the secondary species when exposed to the same climate as that which produced it must revert to the primary, he forgets that this reversion to the winter form is nothing but a reversion – i.e., a sudden return to a primary form through peculiar laws of inheritance – and by no means a gradual re-acquisition of this primary form under the gradual influence of the primary climate. Reversion to the winter form occurs also through other influences, as, for instance, by high temperature. Reversions of this kind, depending on laws of heredity, certainly happen with those cases of transmutation which do not alternate with the primary form, as in seasonal dimorphism, but which occur continuously. They would, however probably be more quickly suppressed in such cases than in seasonal dimorphism, where the constant alternation of the primary and secondary forms must always maintain the tendency of the latter to produce the former.

That the above conclusion is correct – that a secondary species, when exposed to the external conditions under the influence of which the primary form originated, does not again revert to the latter – is proved by experience with plants. Botanists53 assure us “that cultivated races which become wild, and are thus brought back to their former conditions of life, do not become changed into the original wild form, but into some new one.”

A second point which appears to me to be elucidated by seasonal dimorphism, is the origin of variability. It has already been prominently shown that secondary forms are for the most part considerably more variable than primary forms. From this it follows that similar external influences either induce different changes in the different individuals of a species, or else change all individuals in the same manner, variability arising only from the unequal time in which the individuals are exposed to the external influence. The latter is undoubtedly the case, as appears from the differences which are shown by the various individuals of a secondary form. These are always only differences of degree and not of kind, as is perhaps most distinctly shown by the very variable A. Prorsa (summer form), in which all the occurring variations differ only by the Levana marking being more or less absent, and, at the same time, by approximating more or less to the pure Prorsa marking; but changes in a totally different direction never occur. It is likewise further evident, as has been mentioned above, that allied species and genera, and even entire families (Pieridæ), are changed by similar external inducing causes in the same manner – or, better, in the same direction.

In accordance with these facts the law may be stated, that, in butterflies at least, all the individuals of a species respond to the same external influences by similar changes, and that, consequently, the changes brought about by climatic influences take a fixed direction, determined by the physical constitution of the species. When, however, new climatic forms of butterflies, in which natural selection is completely excluded, and the nature of the species itself definitely determines the direction of the changes, nevertheless show variability from the very beginning, we may venture to conclude that every transformation of a species generally begins with a fluctuation of its characters. But when we find the primary forms of butterflies always far more constant, this shows that the continued crossing of the individuals of a species to a certain extent balances the fluctuations of form. Both facts taken together confirm the law formerly enunciated by me,54 that in every species a period of variability alternates with one of (relative) constancy – the latter indicating the culmination, and the former the beginning or end, of its development. I here call to mind this law, because the facts which I advanced at that time, viz., Hilgendorf’s history of the phyletic development of the Steinheim fossil shells, having since become somewhat doubtful, one might easily be inclined to go too far in mistrusting them and refuse to give them any weight at all.55

In the essay just indicated I traced the origin of a certain class of local forms to local isolation. I attempted to show that when a species finds itself in an isolated district in a condition (period) of variability, it must there necessarily acquire somewhat deviating characters by being prevented from crossing with the individuals of other regions, or, what comes to the same thing, a local form must originate. This production of local forms results because the different variations which, for the time being, constitute the variability of the species, would always be in a different numerical proportion in the isolated district as compared with other regions; and further, because constancy is produced by the crossing of these (isolated) varieties among themselves; so that the resultant of the various components is (local) variation. If the components are dissimilar the resultant would also be different, and thus, from a theoretical point of view, there seems to me no obstacle in the way of the production of such local forms by the process of ‘amixia.’ I believe that I have further shown that numerous local forms can be conceived to have arisen through this process of preventive crossing, whilst they cannot be explained by the action of climatic influences.

That I do not deny the existence of true climatic forms in admitting this principle of ‘amixia,’ as has been frequently imagined, appears sufficiently from the treatise in question. The question arises, however, whether climatic influences may not also originate forms by ‘amixia’ by making a species variable. It would be difficult at present to decide finally upon this subject. If, however, in all cases a variation in a certain fixed direction occurred through climatic influences, a form could not arise by ‘amixia’ from such a variability, since the components could then produce resultants different only in degree and not in kind. But we are not yet able to extend our researches to such fine distinctions.

As a final, and not unimportant result of these investigations, I may once more insist that dissimilar influences, when they alternatingly affect a long series of originally similar generations in regularly recurring change, only modify the generations concerned, and not intermediate ones. Or, more briefly, cyclically acting causes of change produce cyclically recurring changes: under their influence series of monomorphic generations become formed into a cycle of di- or polymorphic generations.

There is no occasion to return here to the immediate evidence and proof of the foregoing law. In the latter, however, is comprised the question – is not the cycle of generations produced by cyclical heredity ultimately equivalent to Darwin and Haeckel’s homochronic heredity which forms the ontogenetic stages into a cycle? It is possible that from this point, in the future, the nature of the processes of heredity, which are still so obscure, may be penetrated into, and both phenomena traced to the same cause, as can now be only surmised but not clearly perceived.

Finally, the most general, and in so far chief result of these investigations, appears to me to lie in the conclusion, which may be thus formulated: – A species is only caused to change through the influence of changing external conditions of life, this change being in a fixed direction which entirely depends on the physical nature of the varying organism, and is different in different species, or even in the two sexes of the same species.

I am so little disposed to speak in favour of an unknown transforming power that I may here again insist that the transformation of a species only partly depends upon external influences, and partly on the specific constitution of the particular form. I designate this constitution ‘specific,’ inasmuch as it responds to the same inciting cause in a manner different to the constitution of another species. We can generally form a clear conception why this should be the case; for not only is there in another species a different kind of latent vital activity, but each species has also a different developmental history. It must be admitted that, from the earliest period of the formation of an organism, and throughout all its intermediate stages, properties which have become established, such as growth, nutrition, or tendency to development, have been transferred to the species now existing, each of which bears these tendencies in itself to a certain extent. It is these innate tendencies which determine the external and internal appearance of the species at every period of its life, and which, by their reaction to external factors, represent the life of the individual as well as that of the species. Since the sum of these inherited tendencies must vary more or less in every species, not only is the different external appearance of species as well as their physiological and biological diversity thus explained, but it necessarily follows therefrom, that different species must respond differently to those external causes which tend to produce a change in their form.

Now, this last conclusion is equivalent to the statement that every species, through its physical constitution, (in the sense defined) is impressed with certain fixed powers of variation, which are evidently extraordinarily numerous in the case of each species, but are not unlimited; they permit of a wide range for the action of natural selection, but they also limit its functions, since they certainly restrain the course of development, however wide the latter may be. I have elsewhere previously insisted56 that too little is ascribed to the part played by the physical constitution of species in the history of their transformation, when the course of this transformation is attributed entirely to external conditions. Darwin certainly admits the importance of this factor, but only so far as it concerns the individual variation, the nature of which appears to him to depend on the physical constitution of the species. I believe, however, that in this directive influence lies the precise reason why, under the most favourable external circumstances, a bird can never become transformed into a mammal – or, to express myself generally, why, from a given starting-point, the development of a particular species cannot now attain, even under the most favourable external conditions, any desired goal; and why, from this starting-point, given courses of development, even when of considerable latitude, must be restricted, just as a ball rolling down a hill is diverted by a fixed obstacle in a direction determined by the position of the latter, and depending on the direction of motion and the velocity at the moment of being diverted.

In this sense I agree with Askenasy’s “fixed” direction of variation; but not if another new physical force directing variation itself is thereby intended.57 The explanation of the phenomena does not appear to me to require such an admission, and, if unnecessary, it is certainly not legitimate. According to my view, transmutation by purely internal causes is not to be entertained. If we could absolutely suspend the changes of the external conditions of life, existing species would remain stationary. The action of external inciting causes, in the widest sense of the word, is alone able to produce modifications; and even the never-failing “individual variations,” together with the inherited dissimilarity of constitution, appear to me to depend upon unlike external influences, the inherited constitution itself being dissimilar because the individuals have been at all times exposed to somewhat varying external influences.