Studies in the Theory of Descent, Volume I

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 - P. C. Zeller, “Bemerkungen über die auf einer Reise nach Italien und Sicilien gesammelten Schmetterlingsarten.” Isis, 1847, ii. – xii.]

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 Hoffmann[36 - “Isoporien der europäischen Tagfalter.” Stuttgart, 1873.] 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.

V. On Alternation of Generations

Seasonal dimorphism has already been designated by Wallace as alternation of generation,[37 - [Trans. Linn. Soc., vol. xxv. 1865, p. 9. R.M.]] a term which cannot be disputed so long as it is confined to a regular alternation of dissimilar generations. But little is gained by this definition, however, unless it can be proved that both phenomena are due to similar causes, and that they are consequently brought about by analogous processes. The causes of alternation of generation have, until the present time, been scarcely investigated, owing to the want of material. Haeckel alone has quite recently subjected these complicated phenomena generally to a searching investigation, and has arrived at the conclusion that the various forms of metagenesis can be arranged in two series. He distinguishes a progressive and a retrogressive series, comprising under the former those species “which, to a certain extent, are still in a transition stage from monogenesis to amphigenesis (asexual to sexual propagation), and the early progenitors of which, therefore, never exclusively propagated themselves sexually” (Trematoda, Hydromedusæ). Under the other, or retrogressive form of metagenesis, Haeckel includes a “return from amphigenesis to monogenesis,” this being the case with all those species which now manifest a regular alternation from amphigenesis to parthenogenesis (Aphides, Rotatoria, Daphniidæ, Phyllopoda, &c.). Essentially I can but agree entirely with Haeckel. Simply regarding the phenomena of alternation of generation as at present known, it appears to me to be readily admissible that these multiform modes of propagation must have originated in at least two different ways, which can be aptly formulated in the manner suggested by Haeckel.

I will, however, venture to adopt a somewhat different mode of conception, and regard the manner of propagation (whether sexual or asexual) not as the determining, but only as the secondary cause. I will further hazard the separation of the phenomena of alternating generations (in their widest sense) into two main groups according to their origin, designating the cases of one group as true metagenesis and those of the other as heterogenesis.[38 - It is certainly preferable to make use of the expression “metagenesis” in this special sense instead of introducing a new one. As a general designation, comprehending metagenesis and heterogenesis, there will then remain the expression “alternation of generation,” if one does not prefer to say “cyclical propagation.” The latter may be well used in contradistinction to “metamorphosis.”] Metagenesis takes its origin from a phyletic series of dissimilar forms, whilst heterogenesis originates from a phyletic series of similar forms – this series, so far as we can at present judge, always consisting of similar sexual generations. The former would thus nearly coincide with Haeckel’s progressive, and the latter with his retrogressive metagenesis. Metagenesis may further originate in various ways. In the first place, from metamorphosis, as for example, in the propagation of the celebrated Cecidomyia with nursing larvæ. The power which these larvæ possess of propagating themselves asexually has evidently been acquired as a secondary character, as appears from the fact that there are many species of the same genus the larvæ of which do not nurse, these larvæ being themselves undoubted secondary forms produced by the adaptation of this stage of phyletic development to a mode of life widely different from that of the later stages. In the form now possessed by these larvæ they could never have represented the final stage of their ontogeny, neither could they have formerly possessed the power of sexual propagation. The conclusion seems inevitable that metagenesis has here proceeded from metamorphosis; that is to say, one stage of the ontogeny, by acquiring asexual propagation, has changed the originally existing metamorphosis into metagenesis.

Lubbock[39 - Loc. cit. chap. iv.] is undoubtedly correct when, for cases like that just mentioned, he attempts to derive alternation of generations from metamorphosis. But if we exclude heterogenesis there still remain a large number of cases of true metagenesis which cannot be explained from this point of view.

It must be admitted, with Haeckel, that the alternation of generations in the Hydromedusæ and Trematoda does not depend, as in the case of Cecidomyia, upon the larvæ having acquired the power of nursing, but that the inferior stages of these species always possessed this power which they now only preserve. The nursing Trematode larvæ now existing may possibly have been formerly able to propagate themselves also sexually, this mode of propagation having at the present time been transferred to a later phyletic stage. In this case, therefore, metagenesis was not properly produced by metamorphosis, but arose therefrom in the course of the phyletic development, the earlier phyletic stages abandoning the power of sexual reproduction, and preserving the asexual mode of propagation. A third way in which metagenesis might originate is through polymorphosis. When the latter is combined with asexual reproduction, as is especially the case with the Hydrozoa, metagenesis may be derived therefrom. The successive stages of transformation of one and the same physiological individual do not in these cases serve as the point of departure for alternation of generation, but the different contemporary forms living gregariously into which the species has become divided through functional differentiation of the various individuals of the same stock. Individuals are here produced which alone acquire the power of sexual reproduction, and metagenesis is thus brought about, these individuals detaching themselves from the stock on which they originated, while the rest of the individuals remain in combination, and retain the asexual mode of propagation. No sharp distinction can be otherwise drawn between this and the cases previously considered.[40 - The idea that alternation of generation is derived from polymorphism (not the reverse, as usually happens; i.e. polymorphism from alternation of generation) is not new, as I find whilst correcting the final proof. Semper has already expressed it at the conclusion of his interesting memoir, “Über Generationswechsel bei Steinkorallen,” &c. See “Zeitschrift f. wiss. Zool.” vol. xxii. 1872.] The difference consists only in the whole cycle of reproduction being performed by one stock; both classes have the common character that the different phyletic stages never appear in the same individual (metamorphosis), but in the course of further phyletic development metagenesis at the same time arises, i.e. the division of these stages among a succession of individuals. We are therefore able to distinguish this primary metagenesis from the secondary metagenesis arising from metamorphosis.

It is not here my intention to enter into the ultimate causes of metagenesis; in this subject we should only be able to advance by making vague hypotheses. The phenomenon of seasonal dimorphism, with which this work has mainly to deal, is evidently far removed from metagenesis, and it was to make this clear that the foregoing observations were brought forward. The characters common in the origin of metagenesis are to be found, according to the views previously set forth, in the facts that here the faculty of asexual and of sexual reproduction is always distributed among several phyletic stages of development which succeed each other in an ascending series (progressive metagenesis of Haeckel), whereas I find differences only in the fact that the power of asexual propagation may (in metagenesis) be either newly acquired (larva of Cecidomyia) or preserved from previous ages (Hydroida). It seems that in this process sexual reproduction is without exception lost by the earlier, and remains confined solely to the most recent stages.

From the investigations on seasonal dimorphism it appears that a cycle of generations can arise in an entirely different way. In this case a series of generations originally alike are made dissimilar by external influences. This appears to me of the greatest importance, since seasonal dimorphism is without doubt closely related to that mode of reproduction which has hitherto been exclusively designated as heterogenesis, and a knowledge of its mode of origination must therefore throw light on the nature and origin of heterogenesis in general.

In seasonal dimorphism, as I have attempted to show, it is the direct action of climate, and indeed chiefly that of temperature, which brings about the change in some of the generations. Since these generations have been exposed to the alternating influence of the summer and winter temperature a periodical dimorphism has been developed – a regular cycle of dissimilar generations. It has already been asserted that the consecutive generations of a species comport themselves with respect to heredity in a manner precisely similar to that of the ontogenetic stages, and at the same time such succeeding generations point out the parallelism between metamorphosis and heterogenesis. If influences capable of directly or indirectly producing changes operate on any particular stage of development, these changes are always transmitted to the same stage. Upon this metamorphosis depends. In a precisely similar manner changes which operated periodically on certain generations (1, 3, 5, for instance) are transmitted to these generations only, and not to the intermediate ones. Upon this depends heterogenesis. We have just been led to the comprehension of heterogenesis by cyclical heredity, by the fact that a cycle is produced whenever a series of generations exists under regularly alternating influences. In this cycle newly acquired changes, however minute in character at first, are only transmitted to a later, and not to the succeeding generation, appearing only in the one corresponding, i.e. in that generation which exists under similar transforming influences. Nothing can more clearly show the extreme importance which the conditions of life must have upon the formation and further development of species than this fact. At the same time nothing shows better that the action of these conditions is not suddenly and violently exerted, but that it rather takes place by small and slow operations. In these cases the long-continued accumulation of imperceptibly small variations proves to be the magic means by which the forms of the organic world are so powerfully moulded. By the application of even the greatest warmth nobody would be able to change the winter form of A. Levana into the summer form; nevertheless, the summer warmth, acting regularly on the second and third generations of the year, has, in the course of a lengthened period, stamped these two generations with a new form without the first generation being thereby changed. In the same region two different climatic varieties have been produced (just as in the majority of cases climatic varieties occur only in separate regions) which alternate with each other, and thus give rise to a cycle of which each generation propagates itself sexually.

But even if seasonal dimorphism is to be ascribed to heterogenesis, it must by no means be asserted that those cases of cyclical propagation hitherto designated as heterogenesis are completely identical with seasonal dimorphism. Their identity extends only to their origin and manner of development, but not to the mode of operation of the causes which bring about their transformation. Both phenomena have a common mode of origination, arising from similar (monomorphic) sexual generations and course of development, a cycle of generations with gradually diverging characters coming into existence by the action of alternating influences. On the other hand, the nature of the changes by which the secondary differs from the primary generation may be referred to another mode of action of the exciting causes. In seasonal dimorphism the differences between the two generations are much less than in other cases of heterogenesis. These differences are both quantitatively less, and are likewise qualitative, affecting only characters of biological insignificance.[41 - See my essay “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872.] The variations in question are mostly restricted to the marking and colouring of the wings and body, occasionally affecting also the form of the wing, and in a few cases the size of the body (Plebeius Amyntas), whilst the bodily structure – so far at least as my investigations extend – appears to be the same in both generations.[42 - [In the case of monogoneutic species which, by artificial ‘forcing,’ have been made to give two generations in the year, it has generally been found that the reproductive system has been imperfectly developed in the second brood. A minute anatomical investigation of the sexual organs in the two broods of seasonally dimorphic insects would be of great interest, and might lead to important results. R.M.]]

The state of affairs is quite different in the remaining cases of heterogenesis; here the entire structure of the body appears to be more or less changed, and its size is often very different, nearly all the internal organs differing in the two generations. According to Claus,[43 - “Grundzüge der Zoologie.” 2nd ed. Leipzig, 1872. Introduction.] “we can scarcely find any other explanation of the mode of origination of heterogenesis than the gradual and slow advantageous adaptation of the organization to important varying conditions of life” – a judgment in which this author is certainly correct. In all such cases the change does not affect unimportant characters, as it does in butterflies, but parts of biological or physiological value; and we cannot, therefore, consider such changes to have originated through the direct action of altered conditions of life, but indirectly through natural selection or adaptation.

Thus, the difference between seasonal dimorphism and the other known cases of heterogenesis consists in the secondary form in which the species appears in the former originating through the direct action of external conditions, whilst in the latter this form most probably originates through the indirect action of such influences. The first half of the foregoing proposition is alone capable of provisional proof, but it is in the highest degree probable that the latter half is also correct. Naturally we cannot say to what extent the direct action of external conditions plays also a part in true heterogenesis, as there have been as yet no experiments made on its origin. That direct action, working to a certain extent co-operatively, plays only a secondary part, while the chief cause of the change is to be found in adaptation, no one can doubt who keeps in view, for instance, the mode of propagation discovered by Leuckart in Ascaris nigrovenosa. In this worm, the one generation lives free in the water, and the other generation inhabits the lungs of frogs, the two generations differing from one another in size of body and structure of internal organs to an extent only possible with the true Nematoda.

To prevent possible misunderstanding, let it be finally noted – even if superfluous – that the changes causing the diversity of the two generations in seasonal dimorphism and heterogenesis are not of such a nature that the value of different “specific characters” can be attached to them. Distinctly defined specific characters are well known not to occur generally, and it would therefore be erroneous to attach but little value to the differences in seasonal dimorphism because these chiefly consist in the colouring and marking of the wings. The question here under consideration is not whether two animal forms have the value of species or of mere varieties – a question which can never be decided, since the reply always depends upon individual opinion of the value of the distinctions in question, and the idea of both species and varieties is moreover purely conventional. The question is, rather, whether the distinguishing characters possess an equal constancy – that is, whether they are transmitted with the same force and accuracy to all individuals; and whether they occur, therefore, in such a manner that they can be practically employed as specific characters. With respect to this, it cannot be doubtful for a moment that the colouring and marking of a butterfly possess exactly the same value as the constant characters in any other group of animals, such as the palate-folds in mice, the structure of the teeth in mammals, the number and form of the wing and tail feathers in birds, &c. We have but to remember with what wonderful constancy often the most minute details of marking are transmitted in butterflies. The systematist frequently distinguishes between two nearly allied species, as for instance in the Lycænidæ, chiefly by the position of certain insignificant black spots on the under side of the wing (P. Alexis female, and P. Agestis); and this diagnosis proves sufficient, since P. Alexis, which has the spots in a straight row, has a different caterpillar to P. Agestis, in which the central spot is nearer the base of the hind wing!

For the reasons just given, I maintain that it is neither justifiable nor useful to designate the di- and polymorphism of butterflies as di- and polychroism, and thereby to attribute but little importance to these phenomena.[44 - With reference to this subject, see the discussion by the Belgian Entomological Society, Brussels, 1873.] This designation would be only justifiable if the differences of colour were due to other causes than the differences of form, using this last word in a narrow sense. But it has been shown that the same direct action of climate which originates new colours, produces also in some species differences of form (contour of wing, size, &c.); whilst, on the other hand, it has long been known that many protective colours can only be explained by the indirect action of external conditions.

When I raise a distinction in the nature of the changes between seasonal dimorphism and the remaining known cases of heterogenesis, this must be taken as referring only to the biological or physiological result of the change in the transformed organism itself. In seasonal dimorphism only insignificant characters become prominently changed, characters which are without importance for the welfare of the species; while in true heterogenesis we are compelled to admit that useful changes, or adaptations, have occurred.

Heterogenesis may thus be defined either in accordance with my proposal or in the manner hitherto adopted, since it may be regarded as more morphological than the cyclical succession of differently formed sexual generations; or, with Claus, as the succession of different sexual generations, “living under different conditions of existence” – a definition which applies in all cases to seasonal dimorphism. Varying conditions of existence, in their widest sense, are the result of the action of different climates; and a case has been made known recently in which it is extremely probable that the climatic differences of the seasons have produced a cycle of generations by influencing the processes of nutrition. This case is quite analogous to that which we have observed in the seasonal dimorphism of butterflies, but with the distinction that the difference between the winter and summer generations does not, at least entirely, consist in the form of the reproductive adult, but almost entirely in its ontogeny – in the mode of its development. A comparison of this case with the analogous phenomenon in butterflies, may be of interest. In the remarkable fresh-water Daphnid, Leptodora hyalina Lillejeborg, it was proved some years ago by P. E. Müller,[45 - P. E. Müller, “Bidrag til Cladocerners Fortplantingshistorie,” 1868.] who studied the ontogeny, that this last was direct, since the embryo, before leaving the egg, already possesses the form, members, and internal organs of the adult. This was, at least, the case with the summer eggs. It was subsequently shown by Sars[46 - Sars, in “Förhandlinger i Videnskabs Selskabet i Christiania,” 1873, part i.] that this mode of development only holds good for the summer brood, the winter eggs producing an embryo in the spring which possesses only the three first pairs of limbs, and, instead of compound eyes, only a single frontal eye, thus exhibiting briefly, at first, the structure of a Nauplius, and gradually acquiring that of Leptodora. The mature form derived from the winter eggs is not distinguishable from the later generations, except by the presence of the simple larval eye, which appears as a small black spot. The generations when fully developed are thus distinguished only by this minute marking, but the summer generation undergoes direct development, whilst the winter generation, on the contrary, is only developed by metamorphosis, beginning with the simplest Crustacean type, and thus fairly representing the phyletic development of the species. We therefore see, in this case, the combination of a metamorphic and a direct development taking place to a certain extent under our eyes. It cannot be proved with certainty what the cause of this phenomenon may be, but the conjecture is almost unavoidable that it is closely related to the origin of the seasonal dimorphism of butterflies, since both depend on the alternating climatic influences of summer and winter: it is most probable that these influences have directly[47 - [Eng. ed. Recent researches on alternation of generation in the Daphniacea have convinced me that direct action of external conditions does not in these cases come into consideration, but only indirect action.]] brought about a shortening of the period of development in summer. Thus we have here a case of heterogenesis nearly related to the seasonal dimorphism of butterflies in a twofold manner – first, because the cycle of generations is also in this case brought about by the direct action of the external conditions of life; and secondly, the winter form is here also the primary, and the summer form the secondary one.

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 - See my memoir, “Über Bau und Lebenserscheinungen der Leptodora hyalina,” Zeitschrift f. wiss. Zool., vol. xxiv. part 3, 1874.] 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 Heyden[49 - Stettin. entom. Zeit., vol. xviii. p. 83, 1857.] some years ago, in the case of Lachnus Querci, and has been since confirmed by Balbiani.[50 - Compt. Rend., vol. lxxvii. p. 1164, 1873.]

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 - [“Accidental” in the sense of our being in ignorance of the laws of variation, as so frequently insisted upon by Darwin. R.M.]] 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 - [Eng. ed. Since this was written I have studied the ornamental colours of the Daphniidæ; and, as a result, I no longer doubt that sexual selection plays a very important part in the marking and colouring of butterflies. I by no means exclude both transforming factors, however; it is quite conceivable, on the contrary, that a change produced directly by climate may be still further increased by sexual selection. The above given case of Polyommatus Phlæas may perhaps be explained in this manner. That sexual selection plays a part in butterflies, is proved above all by the odoriferous scales and tufts of the males discovered by Fritz Müller.] [For remarks on the odours emitted by butterflies and moths, see Fritz Müller in “Jena. Zeit. f. Naturwissen.,” vol. xi. p. 99; also “Notes on Brazilian Entomology,” Trans. Ent. Soc. 1878, p. 211. The odoriferous organs of the female Heliconinæ are fully described in a paper in “Zeit. f. Wissen. Zool.,” vol. xxx. p. 167. The position of the scent-tufts in the sphinx-moths is shown in Proc. Entom. Soc. 1878, p. ii. Many British moths, such as Phlogophora meticulosa, Cosmia trapezina, &c. &c., have tufts in a similar position. The fans on the feet of Acidalia bisetata, Herminia barbalis, H. tarsipennalis, &c., are also probably scent organs. A large moth from Jamaica, well known to possess a powerful odour when alive (Erebus odorus Linn.), has great scent-tufts on the hind legs. For the application of the theory of sexual selection to butterflies, see, in addition, to Darwin’s “Descent of Man,” Fritz Müller in “Kosmos,” vol. ii. p. 42; also for January, 1879, p. 285; and Darwin in “Nature,” vol. xxi. January 8th, 1880, p. 237. R.M.]]

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. Botanists[53 - Nägeli, “Entstehung und Begriff der naturhistorischen Art,” Munich, 1865, p. 25. The author interprets the facts above quoted in a quite opposite sense, but this is obviously erroneous.] 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 - See my essay, “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872.] 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 - [Eng. ed. In the summer of 1877, Dr. Hilgendorf again investigated the Steinheim fossil shells, and found his former statements to be completely confirmed. At the meeting of the German Naturalists and Physicists at Munich, in 1877, he exhibited numerous preparations, which left no doubt that the chief results of his first research were correct, and that there have been deposited a series of successively derived species together with their connecting intermediate forms.]]

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 insisted[56 - See my essay, “Über die Berechtigung der Darwin’schen Theorie.” Leipzig, 1868.] 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 - I expressly insist upon this here, because the notice of Askenasy’s thoughtful essay which I gave in the “Archiv für Anthropologie” (1873) has frequently been misunderstood.] 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.