Studies in the Theory of Descent (Volumes 1 and 2)

by August Weismann

STUDIES IN THE THEORY OF DESCENT.

Part III.
ON THE FINAL CAUSES OF TRANSFORMATION.

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III.
THE TRANSFORMATION OF THE MEXICAN AXOLOTL INTO AMBLYSTOMA.

INTRODUCTION.

Since the time when Duméril made known the transformation of a number of Axolotls into the so-called Amblystoma form, this Mexican Amphibian has been bred in many European aquaria, chiefly with the view to establish the conditions under which this transformation occurred, so as to be enabled to draw further conclusions as to the true causes of this exceptional and enigmatical metamorphosis.

Although the Amphibians propagated freely, the cases in which transformation occurred remained extremely rare, and it was not once possible to reply to the main question, viz. whether this metamorphosis was determined by external conditions or by purely internal causes; to say nothing of the possibility of there perhaps being discoverable certain definite external influences by means of which the metamorphosis could have been induced with certainty. But while these points are undecided all attempted theoretical interpretations of the phenomenon must be devoid of a solid basis.

It appeared to me from the first that the history of this transformation of the Axolotl was of special theoretical value; indeed I believed that it might possibly furnish a special case for deciding the truth of those ground-principles, according to which the origin of this species is represented by the two conflicting schools as a case of transformation or as one of heterogenesis. I therefore determined to make some experiments with the Axolotl myself, in the hopes of being fortunate enough to be able to throw some light upon the subject.

In the year 1872 Prof. v. Kölliker was so good as to leave with me five specimens of his Axolotls, bred in Würzburg, and these furnished a numerous progeny in the following year. With these I carried out the idea, the theoretical bearing of which will be shown subsequently, whether it would not be possible to force all the larvæ, or at any rate, the greater majority, to undergo transformation by exposing them to conditions of life which made the use of gills difficult, and that of lungs more easy; in other words, by compelling them to live partly on land at a certain stage of life.

During that year indeed I obtained no results, most of the larvæ perishing before the time for such an experiment had arrived, and the few survivors did not undergo transformation, but lived on to the following spring and then also died one after the other. Through long absence from Freiburg, necessitated by other labours, I had evidently left them without sufficient care and attention. I was thus led to the conviction, which was more fully confirmed subsequently, that no results can be obtained without the greatest care and attention in rearing, towards which single object all one’s interest should be concentrated, and it must not be considered irksome to have to devote daily for many months a large amount of time to this experiment. As it was evident that I could not afford this time without calling in other aid, I hailed with pleasure an opportunity of witnessing the experiment performed by other hands.

A lady living here (Freiburg), Fräulein v. Chauvin, undertook to rear a number of my larvæ of the following year which had just hatched, and in accordance with my idea to make the experiment of forcibly compelling them to adopt the Amblystoma form. How completely this was accomplished will be seen from the following notes by the lady herself, and it will no less appear that these results were only obtained by that care in treatment and delicacy of observation which she devoted to the experiments.

EXPERIMENTS.

“I began the experiments on June 12th, 1874, with five larvæ about eight days old, these being the only survivors out of twelve. Owing to the extraordinary delicacy of these creatures, the quality and temperature of the water, and the nature and quantity of their food exerts the greatest influence, especially in early life, and one cannot be too cautious in their treatment.

“The specimens were kept in a glass globe of about thirty centimeters in diameter, the temperature of the water being regulated; as food at first Daphnids, and afterwards larger aquatic animals were introduced in large numbers. By this means all the five larvæ throve excellently. At the end of June the rudiments of the front legs appeared in the most vigorous specimens, and on the 9th of July the hind legs also became visible. At the end of November I noticed that one Axolotl remained constantly at the surface of the water, and this led me to suppose that the right period had now arrived for effecting the transformation into Amblystoma. For brevity I shall designate this as No. I., and the succeeding specimens by corresponding Roman numerals.

“In order to bring about this metamorphosis, on December 1st, 1874, No. I. was placed in a large-sized glass vessel containing earth arranged in such a manner that, when the vessel was filled with water, only one portion of the surface of the earth was entirely covered by the liquid, and the creature in the course of its frequent perigrinations was thus more or less exposed to the air. The water was gradually diminished on the following days, during which period the first changes made their appearance in the Amphibian—the gills commenced to shrivel up, and at the same time the creature showed a tendency to seek the shallowest spots. On December 4th, it took entirely to the land, and concealed itself among some damp moss which I had placed on a heap of sand on the highest portion of the earth in the glass vessel. At this period the first ecdysis occurred. Within the four days from the 1st to the 4th of December, a striking change took place in the external appearance of No. I., the gill-tufts shrivelled up almost entirely, the dorsal crest completely disappeared, and the tail, which had hitherto been broad, became rounded and similarly formed to that of a land salamander. The grey-brown colour of the body changed gradually into a blackish hue; isolated spots, at first of a dull white, made their appearance and these in time increased in intensity.

“When the Axolotl left the water on December 4th the gill-clefts were still open, but these closed gradually, and after about eight days were overgrown with skin and no longer to be seen.

“Of the other larvæ three appeared at the end of November (i.e. at the same time when No. I. came to the surface of the water) to have kept pace in development with No. I., an indication that for these also the right period had arrived for accelerating the developmental processes. They were therefore submitted to the same treatment as No. I. No. II. became transformed at the same time and exactly in the same manner as the latter; its gill-tufts were complete when it was first placed in the shallow water, but after four days these had almost entirely disappeared; in the course of about ten days after it took to the land, the overgrowth of skin on the gill-clefts and the complete assumption of the salamander form occurred. During this last period the creature took food, but only when urged to do so.

“In Nos. III. and IV. the development proceeded more slowly. Neither of these so frequently sought the shallow spots, nor did they as a rule remain so long exposed to the air, so that the greater part of January had expired before they took entirely to the land. Nevertheless the dessication of the gill-tufts did not take a longer time than in Nos. I. and II. as the first ecdysis occurred as soon as they took to the land.

“No. V. showed still more striking deviations in its transformation than Nos. III. and IV., but as this specimen appeared much weaker than the others from the beginning and was retarded in growth to a most notable extent, this is by no means surprising. It took fourteen instead of four days before the transformation had advanced far enough to enable it to leave the water. It was especially interesting to observe the behaviour of this specimen during this period. Its weak and delicate constitution evidently made it much more susceptible to all external influences than the others. If exposed to the air for too long a time it acquired a light colour, and when annoyed or alarmed it emitted a peculiar odour, similar to that of a salamander. As soon as these phenomena were observed it was at once placed in deeper water, into which it immediately plunged and gradually recovered itself, the gills always becoming again expanded. The same experiment was repeated several times and always led to the same result, from which we may venture to conclude that by accelerating the transformation too energetically, the process may come to a standstill, and even by continued compulsion may end in death.

“It yet remains to be mentioned with respect to Axolotl No. V. that this specimen, unlike all the others, did not emerge from the water at the first ecdysis, but at the time of the fourth.

“All the Axolotls are now (July, 1875) living, and are healthy and vigorous, so that with respect to their state of nourishment there is nothing to prevent their propagating. Of the first four the largest is fifteen centim. long; Axolotl No. V. measures twelve centim.

“The preceding statements appear to demonstrate the correctness of the views advanced in the Introduction:—Axolotl larvæ generally but not always complete their metamorphosis if, in the first place, they emerge sound from the egg and are properly fed; and if, in the next place, they are submitted to the necessary treatment for changing aquatic into aërial respiration. It is obvious that this treatment must only be applied very gradually, and in such a manner as not to overtax the vital energy of the Amphibian.”

* * * * *

To the foregoing remarks of Fräulein v. Chauvin I may add that in all five cases the transformation was complete, and not to be confounded with that change which occurs more or less in all Axolotls in the course of time when confined in small glass vessels. In this last case there frequently appear changes in the direction of the Amblystoma form without the latter being actually reached. In the five adult Axolotls which I possessed for a short time, and of which two were at least four years old, the gills were much shrivelled, but the aquatic tail and dorsal crest were unchanged. The crest may, however, also disappear, and the tail become shortened without these changes being due to a transformation into Amblystoma, as will be shown further on.

With respect to the duration of the transformation, this amounted in Axolotls Nos. I. to IV. altogether to twelve or fourteen days. Of these, four days were taken by the first changes which occurred while the creature was still in the water; the remaining time, to the completion of the metamorphosis, was passed on land. Duméril gives the duration of the metamorphosis as sixteen days.

The following results of the experiments just described appear to me to be especially noteworthy:—The five Axolotl larvæ which can alone be taken into consideration, the others having soon perished, all experienced metamorphosis, and without an exception became Amblystomas. Only one of them, No. I., by persistently swimming at the surface, as was observed at the end of six months, showed a decided tendency to undergo metamorphosis and to adopt aërial respiration. With respect to this specimen it may therefore be confidently assumed that it would have taken to the land, and that metamorphosis would have occurred without artificial aid, just as was the case in the thirty specimens which Duméril altogether observed.

Respecting Nos. II., III., and IV., on the other hand, such a supposition is but little probable. These three larvæ endeavoured to keep in deep water and avoided as long as possible the shallow places which would have enforced them to take entirely to lung breathing. Metamorphosis thus occurred more than a month later in these individuals.

Finally, there can scarcely be any doubt that No. V. would not have become transformed without forcible adaptation to an aërial life.

From these results we may venture to conclude that most Axolotl larvæ change into the Amblystoma form when, at the age of six to nine months, they are placed in such shallow water that they are compelled to respire chiefly by their lungs. The experiments before us are certainly at present but very few in number, but such a conclusion cannot be termed premature if we consider that out of several hundred Axolotls (the exact number is not given) Duméril obtained only about thirty Amblystomas, while v. Kölliker bred only one Amblystoma out of a hundred Axolotls.

It now only remains questionable whether each larva could have been forced to undergo metamorphosis, but this could only be decided by new experiments. It was originally my intention to have delayed the publication of the experiments till Fräulein v. Chauvin had repeated them in larger numbers, but as my Axolotls have not bred this year (1875) I must abandon my scheme, and this can be done the more readily because, for the theoretical consideration of the facts, it is immaterial whether all or only nearly all the Axolotls could have been compelled to undergo transformation. I must not, however, omit to mention that Herr Gehrig, the curator of our Zoological Museum, bred a considerable number of larvæ from the same brood as that with which Fräulein v. Chauvin experimented, and that of these larvæ six lived over the winter without undergoing metamorphosis. They were always kept in deep water and thus furnished the converse experiment to those recorded above; they further prove that this whole brood did not have a previous tendency to undergo metamorphosis.

If these new facts are to be made use of to explain the nature of this extraordinary process of transformation in accordance with our present conception, the data already known must in the first place be called to our aid.

It has first to be established that Siredon Mexicanus never, as far as we know, undergoes metamorphosis in its native country. This Amphibian is there only known in the Siredon form, a statement which I have taken from De Saussure,[225] who has himself observed the Axolotl in the Mexican lakes. This naturalist never found a single Amblystoma in the neighbourhood of the lakes, “nevertheless the larva (Axolotl) is so common there that it is brought into the market by thousands.” De Saussure believes that in Mexico the Axolotl does not undergo transformation.[226] The same statement is distinctly made by Cope,[227] whose specimens of Siredon Mexicanus bred in America, even in captivity showed “no tendency to become metamorphosed.” On the other hand Tegetmeier observed[228] that one out of five specimens obtained from the Lake of Mexico underwent metamorphosis, and this accordingly establishes the second fact, viz. that the true Axolotl becomes transformed under certain conditions into an Amblystoma when in captivity.

This last remark would be superfluous if, as was for a long time believed, the Paris Axolotls, of which the metamorphosis was first observed and which at the time made such a sensation, were actually Siredon Mexicanus, i.e. the Siredon which alone in its native country bears the name of Axolotl. In his first communication Duméril was himself of this opinion; he then termed the animal “Siredon Mexicanus vel Humboldtii,”[229] but subsequently, in his amplified work[230] on the transformation of the Axolotl observed in the Jardin des Plantes, he retracted this view, and after a critical comparison of the five described species of Siredon, he came to the conclusion that the species in the possession of the Paris Museum was probably Siredon Lichenoides (Baird). All the transformations of Axolotls observed in Europe must consequently be referred to this species, since they were—at least as far as I know—all derived from the Paris colony. My own experimental specimens were also indirectly descended from these.

Now it must be admitted that this does not coincide with the fact that the Amblystoma form which Duméril first obtained from his Axolotls agreed with Cope’s species, A. Tigrinum, while on the other hand we learn from Marsh[231] that Siredon Lichenoides (Baird), when it does undergo metamorphosis, becomes transformed into Amblystoma Mavortium (Baird).

Marsh found Siredon Lichenoides in mountain lakes (7000 feet above the sea) in the southwest of the United States (Wyoming Territory), and obtained from them, by breeding in aquaria, Amblystoma Mavortium (Baird). He considers it indeed doubtful whether the Amphibian undergoes this transformation in its native habitat, although he certainly states this opinion without rigorous proof on purely theoretical considerations, because, according to his view, “the low temperature is there less favourable.”[232]

If I throw doubt upon this last statement it is simply because Amblystoma Mavortium is found native in many parts of the United States, viz:—in California, New Mexico, Texas, Kansas, Nebraska, and Minnesota. It is indeed by no means inconceivable that in the mountain lakes where Marsh obtained this species, it may behave differently with respect to metamorphosis than in other habitats, and this appears probable from certain observations upon Triton which will be subsequently referred to.

Meanwhile, in the absence of further observations, we must admit that the Paris Axolotls were not Siredon Lichenoides, but some nearly allied and probably new species. But little information is furnished by observing the course of the transformation, although it is at least established that this Axolotl in its native habitat does not undergo metamorphosis or does so as exceptionally as in Europe. Unfortunately in his papers Duméril gives no precise statement respecting the locality of this species imported from “Mexico”—it is probable that he was himself unacquainted with it, so that I can only state on the authority of Cope that Amblystoma has never been brought from south of the provinces of Tamaulipas and Chilhuahua, i.e. south of the Tropic of Cancer.[233]

This last statement, however, gives no certainty to the matter. Of much more importance is the above-mentioned fact, that the true Axolotl of the lakes about the city of Mexico does not, as a rule, become transformed into an Amblystoma in that locality, although this species in certain cases undergoes metamorphosis when in confinement. From this circumstance and from the fact that the Paris Axolotl likewise experienced but a very small percentage of metamorphosis in captivity, we may venture to conclude that this species also, in its native habitat, either does not become transformed at all or does so only exceptionally.

But there is yet another body of facts which come prominently into view on considering the history of the transformations. I refer to the existence of numerous species of Amblystoma in a natural state. In the “Revision of the genera of Salamandridæ,” published some years ago by Strauch,[234] this author, following Cope,[235] gives twenty species of Amblystoma as inhabiting North America. Although some of these species are based on single examples, and consequently, as Strauch justly remarks, “may well have to be reduced in the course of time,” there must nevertheless always remain a large number of species which live and propagate as true Amblystomas, and of which the habitat extends from the latitude of New York to that of New Mexico. There are therefore true species of Siredon which regularly assume the Amblystoma form under their natural conditions of life, and which propagate in this form, while, on the other hand, there are at least two species which, under their existing natural conditions of life, always propagate as Siredon. It is but another mode of expression for the same facts if we say that the Mexican Axolotl and the Paris Siredon—whether this is Lichenoides or some other species—stand at a lower grade of phyletic development than those species of Amblystoma which propagate in the salamander form. No one can raise any objection to this statement, while the alternative view maintained by all authors contains a theory either expressed or implied which is, as I believe, incorrect, viz. that the Mexican Axolotl has remained at an inferior stage of phyletic development.

All zoologists[236] who have expressed an opinion upon the transformation of the Axolotl, and who are not, like the first observer of this fact, embarrassed by Cuvier’s views as to the immutability of species, regard the phenomenon as though a species, which owing to some special conditions had hitherto remained at a low stage of development, had now through some other influences been compelled to advance to a higher stage.

I believed for a long time that the phenomenon could not otherwise be comprehended, so little was I then in a position to bring all the facts into harmony with this view. Thus in the year 1872 I expressed myself as follows[237]:—“Why should not a sudden change in all the conditions of life (transference from Mexico to Paris) have a direct action on the organization of the Axolotl, causing it suddenly to reach a higher stage of development, such as many of its allies have already attained, and which obviously lies in the nature of its organization—a stage which it would perhaps itself have reached, although later, in its native country? Or is it inconceivable that the sudden removal from 8000 feet above the sea (Mexican plateau) to the altitude of Paris, may have given the respiratory organs an impetus in the direction of the transformation imminent? In all probability we have here to do with the direct action of changed conditions of life.”

That the substance of this last statement must still hold good is obvious from the experiments previously described, which show that by the application of definite external influences, we have it to a certain extent in our power to produce the transformation. It is precisely in this last point that there lies the new feature furnished by these experiments.

But are we also compelled to interpret the phenomenon in the above manner? i.e. as a sudden advance in the phyletic development of the species occurring, so to speak, at one stroke? I believe not.

What first made this view appear to me erroneous, was the appearance of the living Amblystomas bred from my Axolotl larvæ. These creatures by no means differed from the Axolotls merely in single characters, but they were distinct from the latter in their entire aspect; they differed in some measure in all their parts, in some but slightly and in other parts strongly—in brief, they had become quite different animals. In accordance with this, their mode of life had become completely modified; they no longer lived in the water, but remained concealed by day among the damp moss of the vivarium, coming forth only by night in search of food in dry places.

I had been able to perceive the great difference between the two stages of development from the anatomical data with which I had long been familiar, and which Duméril had made known with respect to the structure of his Amblystomas. But the collecting of numerous details gives no very vivid picture of the metamorphosis; it was the appearance of the living animal that first made me conscious how deep-seated was the transformation which we have here before us, and that this change not merely affected those parts which would be directly influenced by the change in the conditions of life, such as the gills, but that most if not all the parts of the animal underwent a transformation, which could in part be well explained as morphological adaptation to new conditions of life, and partly as a consequence of this adaptation (correlative changes), but could not possibly be regarded as the sudden action of these changed conditions.

Such at least is my view of the case, according to which a per saltum development of the species of such a kind as must here have taken place, is quite inconceivable.

I may venture to assume that most observers of the metamorphosis of Axolotl have, like myself, not been hitherto aware of the extent of the transformation, and it may thus be explained why the theoretical bearings of the case have on all sides been incorrectly conceived. We have here obviously a quite extraordinary case of the first order of importance. I believe that it can easily be shown that the explanation of the history of the metamorphosis of the Paris Axolotl which has hitherto been pretty generally accepted, necessarily comprises a very far-reaching principle. If this interpretation is correct, then in my opinion must also hold good the ideas of those who, like Kölliker, Askenasy, Nägeli, and, among the philosophers, Hartmann and Hübner, would refer the transformation of species in the first instance to a power innate in the organism, to an active, i.e. a self-urging “law of development”—a phyletic vital force.

Thus, if the Axolotls transformed into Amblystomas are regarded as individuals which, impelled by external influences, have anticipated the phyletic development of the others, then this advance can only be ascribed to a phyletic vital force, since the transformation is sudden, and leaves no time for gradual adaptation in the course of generations. The indirect influence of the external conditions of life, i.e. natural selection, is thus excluded from the beginning. But the direct action of the changed conditions of life by no means furnishes an explanation of the complete transformation of the whole structure, such as I have already alluded to, and which I will now enter into more closely.

The differences between the Paris Axolotl and its Amblystoma according to Duméril, Kölliker, and my own observations are as follow:—

1. The gills disappear; the gill-clefts close up, and of the branchial arches only the foremost remain, the posterior ones disappearing. At the same time the os hyoideum becomes changed (Duméril).

2. The dorsal crest completely disappears (Duméril).

3. The aquatic tail becomes transformed into one like that of the salamanders (Duméril), which, however, is not as in the salamander cylindrical, but somewhat compressed laterally (Weismann).

4. The skin becomes yellowish white, irregularly spotted on the sides and back (Duméril), whilst at the same time its former grey-black ground-colour changes into a shining greenish black (Weismann); it loses, moreover, the slimy secretion of the skin, and the cutaneous glands become insignificant (Kölliker).

5. The eyes become prominent and the pupils narrow (Kölliker), and eyelids capable of completely covering the eyes are formed; in Axolotl only a narrow annular fold surrounds the eyes, so that these cannot be closed (Weismann).

6. The toes become narrowed and lose their skin-like appendages (Kölliker), or more precisely, the half webs which connect the proximal extremities of the toes on all the feet (Weismann).

7. The teeth are disposed in this species, as in all other Amblystomæ, in transverse series; whilst in Axolotl, as in Triton larvæ, they are arranged at the sides of the jaw in the form of a curved arch-like band beset with several rows of teeth.[238] (Duméril. See his fig., loc. cit. p. 279).

8. In Axolotl the lower jaw, in addition to the teeth on the upper edge of the bone, also bears “de très petites dents disposées sur plusieurs rangs;” these last disappear after metamorphosis (Duméril). I will add that the persistent teeth belong to the os dentale of the lower jaw, and those that are shed to the os operculare.[239]

9. The surface of the posterior moveable part of the body is slightly concave both before and after transformation; the anterior part is, however, less concave in Amblystoma than in Siredon (Duméril).

I have not yet been able to verify Duméril’s 7th and 9th statements, as I did not want to kill any of my living Amblystomas,[240] simply in order to confirm the observations of a naturalist in whom one may certainly place complete confidence. Neither have I as yet observed the transformation of the branchial arches, but all the other statements of Kölliker and Duméril I can entirely corroborate.

The structural differences between Axolotl and Amblystoma are considerably greater and of more importance than those between allied genera, or indeed than between the families of the Urodela. The genus Siredon undoubtedly belongs to a different sub-order to the genus Amblystoma into which it occasionally becomes transformed. Strauch, the most recent systematic worker at this group, distinguishes the sub-order Salamandrida from that of the Ichthyodea by the possession of eyelids, and by the situation of the palatine teeth in single rows on the posterior edge of the palatal bone: in Ichthyodea the eyelids are wanting and the palatine teeth are either “situated on the anterior edge of the palatal bone,” or “cover the whole surface of the palatal plates in brush-like tufts.”

How is it possible to regard such widely divergent anatomical characters as changes suddenly produced by the action (but once exerted) of deviating conditions of life? Hand in hand with the shedding of the old and the appearance of new palatine teeth, there occurs a change in the anatomical structure of the vertebral column, and also—as we may fairly conclude from Kölliker’s correct observation of the cessation of the slimy secretion—in the histological structure of the skin. Who would undertake to explain all these profound modifications as the direct and sudden action of certain external influences? And if any one were inclined to explain such changes as a consequence of the disappearance of the gills, i.e. as correlative changes, what else is such a correlation than the phyletic vital force under another name?

If from one change arising from the direct action of external agencies, the whole body can in two days become transformed in all its parts, in the precise manner which appears best adapted for the new conditions of life under which it is henceforward to exist, then the word “correlation” is only a phrase which explains nothing, but which prevents any attempt at a better explanation, and it would be preferable to profess simply the belief in a phyletic vital force.

Moreover, it is hardly permissible to seek such an explanation, since Urodela are known which have no gills in the adult state, and which nevertheless possess all the other characters of the Ichthyodea, viz. want of eyelids, characteristic palatine teeth, and the tongue bone. This is the case with the genera Amphiuma (Linn.), Menopoma (Harl.), and Cryptobranchus (v. d. Hoev.). The two first genera, as is known, still possess gill-clefts, but Cryptobranchus has even lost these clefts, which, as in Amblystoma, are overgrown by skin; nevertheless Cryptobranchus is, according to the concurrent testimony of all systematists, a true salamander in habits, tongue bone, palatine teeth,[241] &c. It must further be added that the Axolotl itself can lose the gills without thereby becoming transformed into an Amblystoma. I have previously mentioned that in Axolotls which were kept in shallow water the gills frequently became diminutive, and it also sometimes happens that they completely shrivel up. I possess an Axolotl preserved in alcohol in which the gills have shrivelled up into small irregular bunches, and the dorsal crest is also so completely absent that its place is occupied by a long furrow, and even on the tail the crest has entirely disappeared from the lower edge and about half from the upper edge. Notwithstanding this, the creature is widely removed from Amblystoma in structure; it possesses the arched branchial apparatus, the palatine teeth, the skin, &c., of the Axolotl.

These facts prove, therefore, that the shedding of the gills by no means always entails all the other modifications which we observe in the metamorphosis of Axolotl, so that these modifications are thus not by any means the necessary and immediate consequence of such gill shedding.

Whether these modifications will occur after a long series of generations—whether the successors of Cryptobranchus will also one day acquire the salamandriform structure is another question, and one which I could not exactly answer in the negative. But this question does not here come into consideration, as we are now only concerned with the immediate result of the shedding of the gills.

The problem appears therefore to be as follows:—Either the hitherto received interpretation of the transformational history of the Axolotl as a further development of the species is incorrect, or else the case of Axolotl incontestably proves the existence of a phyletic vital force.

We have now to ask whether the facts of this transformational history are not capable of another explanation.

I believe that this is certainly possible, and that another interpretation can be shown to be correct with some degree of probability.

I am of opinion that those Amblystomas which have been developed in captivity in certain instances from Siredon Mexicanus (S. Pisciformis), as well as from the Paris Axolotls, are not progressive, but reversion forms; I believe that the Axolotls which now inhabit the Mexican lakes were Amblystomas at a former geological (or better, zoological) epoch, but that owing to changes in their conditions of life, they have reverted to the earlier perennibranchiate stage.

I was undoubtedly first led to this conception by the results which arose from my studies on the seasonal dimorphism of butterflies.[242] In this case we were also concerned with the two different forms under which one and the same species appears, and of which it was shown to be probable that the one is phyletically older than the other. The younger summer form, according to my view, has arisen, through the gradual amelioration of the climate, from the winter form, which at an earlier zoological epoch was the only one in existence; but the latter, the primary form, has not for this reason ceased to exist, but now alternates in each year as a winter form with the secondary summer form.

Now with seasonally dimorphic butterflies, it was easily possible to induce the summer brood to assume the winter form by exposing their pupæ for a long time to a low temperature; and it was shown to be highly probable that this abrupt and often very extensive change or transformation, only apparently takes place suddenly, and is but the apparent result of the action of cold upon this generation, whilst in fact it depends upon reversion to the primary form of the species, so that the low temperature, which is only once applied, gives but the impetus to reversion, and is not the true cause of the transformation. This cause must rather be sought in the long continued action of the cold to which the ancestors of our existing butterflies were subjected for thousands of generations, and of which the final result is the winter form.

If we assume for an instant that my interpretation of the transformation of Axolotl as just offered is correct, we should have conditions in many respects analagous to those of seasonal dimorphism. It is true that in this case the two forms no longer alternate regularly with each other, but the primary form may occasionally appear instead of the secondary form, owing to the action of external conditions.

Just as in the case of seasonal dimorphism it is possible to compel the summer generation to abandon the summer form, and to assume the winter guise by the action of cold; so in the present case we are able to induce the Axolotl to adopt the Amblystoma form by making aërial respiration compulsory at a certain stage of life; and further, just as in seasonal dimorphism it can be shown that this artificially produced change is only apparently an abrupt transformation, and is actually a reversion to the much older winter form; so here we have not an actual, but only an apparent remodelling of the species—a reversion to the phyletically older form.

This certainly appears a paradox, inasmuch as a form here arises by reversion which must yet undoubtedly rank as the more highly developed. I believe, however, that much which seems paradoxical in this statement will disappear on further examination.

It must in the first place be taken into consideration that the phyletic development of species need not by any means always take place by advancement. We have indeed many cases of retrogressive development, although in a somewhat different sense, as with parasites and those forms which have degenerated from free locomotion to a sedentary mode of life.[243] I do not confuse this kind of retrogressive development, arising from the arrest of certain organs and systems of organs, with true reversion. The latter is a return to a form which has already been once in existence; but in the former case, in spite of all simplification of the organization, some entirely new feature always comes into existence. But I am not able to see any absurdity in the assumption that even true reversion, whether of a whole species or of the individuals of a certain district, may be regarded as possible, and I require no further concession. Why, for example, should it be inconceivable that at a very remote period the Axolotl was adapted to a life on land; that through the direct and indirect action of changed conditions of life it gradually acquired the salamander form, but that subsequently, through new and unfavourable changes in the conditions of life, it again relapsed to the older form, or at least to one nearly related thereto?

At any rate such an assumption contains nothing opposed to known facts, but can be supported in many ways, and finally it commends itself, at least in my opinion, as offering the only admissible explanation of the facts before us.

The existence of a whole series of species of Amblystoma, as already mentioned, at once shows that species of Siredon can become elevated into the salamander form, and can propagate regularly in this state, and further, that this phyletic advance has already actually taken place in many species.

That degeneration may also occur from this high stage to a lower stage of development, is shown by many observations on our water-salamanders. It is known that under certain circumstances Tritons, as it is generally expressed, become “sexually mature in the larval condition.”

In the year 1864 De Filippi[244] found fifty Tritons in a pool at Andermatten, in the neighbourhood of Puneigen, and of these only two showed the structure of the adult water-salamander; all the others still possessed gills, but notwithstanding this, they agreed in both sexes, in size and in the development of the sexual organs, with mature animals. De Filippi established that these “sexually mature larvæ” not only resembled larvæ externally through the possession of gills, but that they also possessed all the other anatomical characters of the larvæ, i.e. the characteristic bunches of palatine teeth situated on both sides in the position of the subsequent single rows, and a vertebral column represented throughout its whole length by the chorda dorsalis.

According to my view this would be a case of the reversion of the Triton to the immediately anterior phyletic stage, i.e. to the perennibranchiate stage, and in the present instance the majority of zoologists who take their stand by the theory of descent, would certainly concur in this view. I should at least consider it to be a useless play upon words did we here speak of larval reproduction, and thereby believe that we had explained something. The animal certainly becomes sexually mature in the same condition as that in which it first appears as a larva, but we first get an insight into the nature of this process by considering that this so-called “sexually mature larva” has the precise structure which must have been possessed by the preceding phyletic stage of the species, and that an individual reversion to the older phyletic stage of the species is consequently before us. I maintain that Duméril is in error in regarding this case of the Triton as parallel with the true larval reproduction of Wagner’s Cecidomyia larva. In this last case it is certainly not reversion to an older phyletic stage that confers the power of reproduction upon the larvæ, since the latter do not represent an older phyletic stage of the species, but must have arisen contemporaneously with this last stage. The enormous structural difference between the larvæ and the imagines is not explained by the latter having arisen from the former supplementarily as a finished production, but by both having been contemporaneously adapted to continually diverging conditions of life.[245] Considered phyletically, these larvæ are by no means necessarily transitional to the origination of the flies. They could have been quite different without the form of the imagines having been thereby modified, since the stages of insect metamorphosis vary independently of each other in accordance with the conditions of life to which they are subjected, and exert scarcely any, or only a very small form-determining influence upon each other, as has been amply proved in the preceding essay. In any case the power of these larvæ (the Cecidomyiæ) to propagate themselves asexually was first acquired as a secondary character, as appears from the fact that there exist numerous species of the same genus which do not “nurse.” In the form which they now possess they could never have played the part of the final stage of the ontogeny, nor could they formerly have possessed the power of sexual reproduction.[246] In brief, we are here concerned with true larval reproduction, whilst in Triton we have reversion to an older phyletic stage.[247]

I cannot agree with my friend Professor Haeckel when he occasionally designates the reversion of the Tritons as an “adaptation” to a purely aqueous existence.[248] We could here only speak of “adaptation” if we took the word in a quite different sense to that in which it was first introduced into science by Darwin and Wallace. These naturalists thereby designate a gradual bodily transformation appearing in the course of generations in correspondence with the new requirements of altered conditions of life or, in other words, the action of natural selection, and not the result of a suddenly and direct acting transforming cause exerted but once on a generation.

Just because the word “adaptation” can be used in ordinary language in many senses, it is desirable that it should have only one precise signification, and above all that we should not speak of adaptation where scarcely any morphological change occurs, but only a kind of functional change in the sense used by Dohrn.[249] This is the case for example, when Forel[250] shows that fresh water Pulmonifera, the organization of which is attributed to the direct respiration of air, can nevertheless become settled in the greatest depths of mountain lakes through their lungs being again employed as gills. That not the least change in the lungs hereby takes place is shown by the observations of Von Siebold,[251] who saw the shallow water Pulmonifera using their lungs alternately for direct aërial and aquatic respiration, according to the amount of air contained in the water. If with Von Siebold we merely apply the word “adaptation” to such cases, this expression would lose the special sense which it originally conveyed, and the word would have to be abandoned as a terminus technicus; still, such cases may perhaps be spoken of as physiological adaptation.

In any case the reproductive “larvæ” of the Tritons as little present a case of true adaptation as the Axolotl, which occasionally becomes transformed into an Amblystoma. In both cases the transformation referred to is by no means indispensable to the life of the individual. Mature Tritons (devoid of gills) can exist, as I have myself seen, for many months, and probably also for a year in deep water, although adapted for purely pulmonary respiration; whilst Axolotls, as I have already mentioned, can live well for a year in shallow water poor in air. If their gills by this means become shrivelled up or completely disappear, even this is not adaptation in the Darwinian sense, but the effect of directly acting external influences, and chiefly of diminished use.

A case entirely analagous to that of Filippi’s was observed by Jullien in 1869. Four female larvæ of Lissotriton Punctatus (Bell)—(synonymous with Triton Tæniatus, Schnd.), taken from a pool, proved to be sexually mature. They contained mature eggs in their ovaria ready for laying, and two of them actually deposited eggs. Four male larvæ found in the same pool, appeared to be equally developed with respect to size, but their testicles contained no free spermatozoa, but only sperm-cells.[252]

I have met with a third case of a similar kind mentioned by Leydig in his memoir, rich in interesting details, “on the tailed Amphibians of the Wurtemburg fauna.”[253] Schreibers, the former director of the Vienna Museum, also found “larvæ” of Tritons with well-developed gills, but of the size of the “adult male individuals,” and, as shown by anatomical investigation, with well “developed sexual organs,” the ovaria especially being distended with eggs.

It is thus established that species which long ago reached the salamander stage in phyletic development, may occasionally degenerate to the perennibranchiate stage. This fact obviously makes my conception of the Axolotl as a reversion form appear much less paradoxical—indeed, the cases of reversion in Triton are precisely analagous to the process which I suppose to have taken place in the Axolotl. We have only to substitute Amblystomas for Tritons, to imagine the pool in which De Filippi found his “sexually mature Triton larvæ” enlarged to the size of the Lake of Mexico, and to conceive the unknown, and perhaps here transitory, causes of the reversion to be permanent, and we have all that is necessary, so far as we at present know, for the restoration of the Axolotl; we obtain a perennibranchiate population of the lake.

It has not yet been determined whether the perennibranchiate form of the Triton actually prevailed permanently in De Filippi’s pool, since, so far as I know, this has not since been examined.

Let us, however, assume for an instant that this is really the case, and that there exists at that spot a colony of sexually reproductive perennibranchiate Tritons: should we wonder if a true Triton occasionally appeared among their progeny, or if we were able to induce the majority of the individuals of this brood to become metamorphosed into Tritons by keeping them in shallow water? According to my view this is precisely the case of the Mexican Axolotl.

I need not, however, restrict myself to this in order to support my hypothesis, but must also directly combat the view hitherto received, since the latter is in contradiction with facts.

Did there really exist in the Axolotl a tendency to sudden phyletic advancement, then one fact would remain quite incomprehensible, viz. the sterility of the Amblystomas.

Out of about thirty Amblystomas obtained by Duméril down to the year 1870, there was not one in a state of sexual maturity; neither copulation nor deposition of eggs took place, and the anatomical investigation of single specimens showed that the eggs were immature, and that the spermatozoa, although present, were without the undulating membrane characteristic of the salamanders, but were not devoid of all power of movement, only, as established by Quatrefages, were “incompletely motile.”[254]

So also the five Amblystomas about which I have been writing, show up to the present time no appearance of reproduction.

The objection raised by Sacc,[255] that the sterility of the Amblystomas bred from Axolotls is attributable to “bad nourishment,” is obviously of but little avail. How is it that the Axolotls, which are fed in a precisely similar manner, propagate so readily? Moreover, I am able to expressly assert that my Amblystomas were very well fed. It is true that they have as yet scarcely reached the age of two years, but the Axolotl propagates freely in the second year, and some of Duméril’s Amblystomas were five years old in 1870.

This fact of the sterility is strongly opposed to the idea that these Amblystomas are the regular precursors of the phyletically advancing genus Siredon.[256] I will by no means assert that my theory of reversion actually explains the sterility, but it is at least not directly opposed to it. Mere reversion forms may die off without propagating themselves; but a new form called forth by the action of a phyletic vital force should not be sterile, because this is the precise “aim” which the vital force had in view. The conception of a vital force comprises that of teleology.

The sterility of Amblystoma moreover, although not completely explicable from our standpoint, can be shown to be a phenomenon not entirely isolated. In the above mentioned case of Lissotriton Punctatus, the female “larvæ” were certainly sexually mature and laid eggs, but the males of the same period contained in their testicles no fully developed spermatozoa.

Other cases of this kind are unknown to me; at the time when I made the experiments with butterflies already recorded (see the first essay), this point of view was remote, and I therefore neglected to examine the artificially bred reversion forms with respect to their organs of reproduction. But general considerations lead to the supposition that atavistic forms may easily remain sterile.

Darwin[257] finds the proximate causes of sterility in the first place in the action of widely diverging conditions of life, and in the next place in the crossing of individuals widely different in constitution. Now it is certainly deviating conditions of life which lead to the metamorphosis of the Axolotl, and from this point of view it cannot be surprising if we find those individuals sterile which show themselves so especially affected by these changed conditions as to revert to the salamander form.

By this it is not in any way meant to be asserted that reversion is invariably accompanied by sterility, and one cannot raise as an objection to my interpretation of the metamorphosis of the Axolotl, that a reproductive colony of Axolotls could never have arisen by reversion. On the contrary, Jullien’s egg-depositing female Triton larvæ show that also with reversion the power of reproduction may be completely preserved.[258] From the above-mentioned general causes of sterility, it may even be inferred that fertility can be lost in different degrees, and it can be further understood to a certain extent why this fertility is more completely lost by reversion to the Amblystoma, than by the reversion of the Triton to the perennibranchiate form.

If in these cases the reversion is brought about by a change in the conditions of life, we may perhaps suppose that the magnitude of this change would determine the degree of fertility, and the preservation of the reversion form. Still more, however, would the fertility be influenced by the extent of the morphological difference resulting from the reversion. We know that the blending of very different constitutions (e.g. the crossing of different species) produces sterility. Something similar results from the sudden reversion to a stage of development widely different in its whole structure. Here also we have in a certain sense the union of two very different constitutions in one individual—a kind of crossing.

From this point of view it can in some measure be comprehended why sterility may be a result of reversion; on the other hand, we thereby obtain no explanation why, with the same amount of morphological difference, in one case complete sterility, and in another relative fertility occurs. The morphological difference between Axolotl and Amblystoma is exactly the same as between Triton and its “sexually mature larva;” the difference between the two cases of reversion depends entirely upon the direction of the leap, that taken in the former case being precisely opposite in direction to that taken in the latter.

Herein might be sought the explanation of the different strength with which the reproductive power is affected; not indeed in the direction of the leap itself, but in the differences in the ontogeny which are determined by the differences in the direction of the leap. The reversion of the Triton to an older phyletic stage coincides with the arrest at a younger ontogenetic stage; or, in other words, the older stage of the phylogeny to which reversion takes place is still entirely comprised in the ontogeny of each individual. Each Triton is perennibranchiate throughout a long period of its life; the reverting individual simply reverts to the older phyletic stage by remaining at the larval stage of its individual development.

But it is quite different with the reversion of the Axolotl to the formerly acquired, but long since abandoned Amblystoma form. This is not retained in the ontogeny of Axolotl, but has been completely lost; for a long series of generations—so must we suppose—the ontogeny has always only attained to the perennibranchiate form. Now if at the present time certain individuals were compelled to revert to the Amblystoma form, certainly no greater leap would have been made from a morphological point of view, than in the reversion of Triton to the perennibranchiate form, but at the same time the leap would be in another direction, viz. over a long series of generations back to a form which the species had not produced for a long period, and which had to a certain extent become foreign to it. We should thus have here also the grafting of a widely different constitution upon that of the Axolotl, or, if one prefers it, the commingling of two widely different constitutions.

Of course I am far from wishing to pretend that this “explanation” is exact; it is nothing more than an attempt to point out the direction in which the causes affecting the reproductive powers in different degrees are to be looked for. A deeper penetration into and special demonstration of the manner in which these causes bring about such results, must be reserved for a future period. For the present it must suffice to have indicated that there is an essential distinction between the two kinds of reversion, and to have made it to some extent comprehensible that this distinction may be the determining impulse with respect to the question of sterility. Perhaps the law here concealed from us may one day be thus formulated:—Atavistic individuals lose the power of reproduction the more completely, the greater the number of generations of their ancestors whose ontogeny no longer comprises the phyletically older stage to which the reversion takes place.

The hypothesis which interprets the transformation of the Axolotl as a case of reversion, thus holds out the possibility of our being able to comprehend the sterility of the Amblystomas arising in this manner, whilst, on the other hand, for the adherents of a phyletic vital force, not only is this observed sterility as Duméril expresses it “un véritable énigme scientifique,” but an absolute paradox. We should expect such a directive and inciting principle to call into existence new forms having vitality and not destined to perish, the more so when it is concerned with a combination of structural characters which, when originating in another manner (viz. from other species of Siredon), have long since shown themselves to have vitality and reproductive power. We are indeed acquainted with species of Amblystoma which propagate as such, and each of which arises from an Axolotl-like larva. Thus we cannot regard the sterile Amblystomas produced by the Paris Axolotls as abortive attempts of a vital force—an interpretation which is certainly in itself already sufficiently rash.

Now if it be asked what change in the conditions of life could have led to the reversion in the Lake of Mexico[259] of the Amblystoma to the Siredon form, I must admit that I can only offer a conjectural reply, having but a conditional value so long as it is not supported by a precise knowledge of the conditions there obtaining, and of the habits both of the Axolotl and of the Amblystoma.

It may be supposed generally that reversion is brought about by the same external conditions as those which formerly produced the perennibranchiate stage. This supposition is in the first place supported by the experiments here recorded, since it is evidently the inducement to aërial respiration which causes the young Axolotl to revert to the Amblystoma form, i.e. the inciting cause under whose domineering influence the Amblystoma form must have arisen.

Here again the case is quite similar to that of seasonally dimorphic butterflies. Reversion of the summer brood to the winter form is there most easily caused by the action of cold, i.e. by the same influence as that under whose sway the winter form was developed.

We know indeed that reversion may also arise by the crossing of races and species, and I have attempted to show that reversion in butterflies may also be brought about by other influences than cold; but still the most probable supposition obviously is, that reversion would be caused by the persistent action of the same influences as those which in a certain sense created the perennibranchiate form. That the latter was produced under the influence of an aquatic life there can be no doubt, and thus, in accordance with my supposition, the hypothetical Amblystoma Mexicanum, the supposed ancestral form of the Axolotl of the Mexican Lake, might have been caused to revert to the perennibranchiate form by a reduction in the possibilities of its living upon land, and by its being compelled to frequent the water.

I will not here return to the consideration of every other opinion ab initio. It is very advisable to distinguish between the mere impulses which are able to produce sudden reversion, and between actual transforming causes which result directly or indirectly in the remodelling of a species. Thus, it is conceivable à priori that reversion may occur by the action of an inciting cause having nothing to do with the origin of the phyletically older form. Temperature can certainly have played no part, or only a very small part, in the formation of the perennibranchiate form; nevertheless cold may well have been one of the inciting causes which induced the Amblystoma at one time to revert to the Siredon form, and we cannot at present consider De Saussure to be incorrect when he maintains that the low temperature of the Mexican winter might prevent that transformation (of the Axolotl into the Amblystoma) which would occur “in the warm reptile-house” of the Jardin des Plantes. He supports this view by stating that “Tschudi has found the Amblystoma” (of course another species) “in the hottest parts of the United States.” “On the Mexican plateau, however, it snows every winter, and if the lake does not actually freeze, its temperature must fall very considerably in the shallowest parts.”

But although this view is not opposed by any theoretical considerations, I still hold it to be incorrect. I doubt whether it is temperature that has brought about the reverse transformation of the Amblystoma into the Axolotl, or which, according to De Saussure’s conception, at the present time prevents the transformation of the Axolotl in the Lake of Mexico. I doubt this because Amblystomas are now known from all parts of the United States as far north as New York, a proof that a winter cold considerably greater than that of the Mexican plateau is no hindrance to the metamorphosis of the Axolotl, and that the genus does not show itself to be in this respect more sensitive than our native genera of Salamandridæ.

The following observations of De Saussure, in which he calls attention to the nature of the Mexican Lake, appear to me to be more worthy of consideration:—“The bottom of this lake is shallow, and one passes imperceptibly from the lake into extensive marshy regions before reaching solid ground; perhaps this circumstance makes the Axolotl incapable of reaching dry land, and prevents the transformation.”

In any case the Lake of Mexico offers very peculiar conditions for Amphibian life. My esteemed friend Dr. v. Frantzius has called my attention to the fact that this lake—as well as many other Mexican lakes—is slightly saline. At the time of the conquest of Mexico by Ferdinand Cortez, this circumstance led to the final surrender of the city, as the Spaniards cut off the supply of water to the besieged, and the water of the lake is undrinkable. The ancient Mexicans had laid down water-conduits from the distant mountains, and the city is still supplied with water brought through conduits.

Now this saltness cannot in itself be the cause of the degeneration to the perennibranchiate form, but it may well be so in combination with other pecularities of the lake. The narrowest part of the lake is the eastern, and it is only in this part that the Axolotl lives. Now in winter, violent easterly gales rush down from the mountains and blow continuously, driving the water before them to such an extent that it becomes heaped up in the western portion of the lake, where it frequently causes floods, whilst 2000 feet of the shallow eastern shore are often laid completely dry.[260]

Now if we consider these two peculiarities, viz. salineness and periodical drying up of a part of the bottom of the lake through continuous gales, we certainly have for the Axolotl, conditions of life which are only to be found in few species. One might certainly attempt to apply these facts in a quite opposite sense, and to regard them as unfavourable to my theory, since the retreat of the water from a great portion of the bottom of the lake would—so one might think—rather facilitate transition to a life upon land, and indeed compel the adoption of such a mode of existence. But we should thus forget that the exposed bottom of the lake is a sterile surface without food or place of concealment, and, above all, without vegetation; and further, that owing to the considerable salineness of the water (specific gravity = 1.0215),[261] the whole of the exposed surface must be incrusted with salt, a circumstance which would render it quite impossible for the creatures to feed upon land. Sodic chloride and carbonate are dissolved in the water in such considerable quantities, that they are regularly deposited upon the shores of the lake as a crust, which is collected during the dry season of the year and sent into the market under the name of “tequisquite” (Mühlenpfordt).[262]

Thus the supposition is not wanting in support, that peculiar conditions make it more difficult for the creature to obtain its food upon land than in the water, and this alone may have been sufficient to have induced it to acquire the habits of a purely aquatic existence, and thus to revert to the perennibranchiate or Ichthyodeous form.

But enough of supposition. We must not complain that we are unable from afar to discover with precision the causes which compelled the Axolotl to abandon the Amblystoma stage, as long as we are not able to explain the much nearer cases of reversion in Filippi’s and Jullien’s Tritons; nevertheless, in these cases also, the causes affecting the whole colony of Tritons must be general, since—at least in the case noticed by Filippi—the greater majority of the individuals remained in the larval condition. Experiments with Triton larvæ could throw greater light upon this subject; it would have in the first place to be established whether reversion could be artificially induced, and if so, by what influences.

From the previously mentioned experiments with butterflies, as well as from the results obtained with Axolotls, we should expect that in Tritons, reversion to the Ichthyodeous form would take place if we allowed the inciting cause, viz. the bathing of the gills and of the whole body with water, to act persistently, and at the same time withheld that influence under whose action the salamander form became developed, viz. the bathing of the gills, the skin, and the surfaces of the lungs with air.

Old experiments of this kind are to be met with, but they were never carried on for a sufficient time to entirely allay the suspicion, that the specimens concerned would perhaps have undergone the ordinary metamorphosis if their existence had been prolonged.

Thus, Schreibers[263] relates that “by confining tadpoles of the salamander found at large in their last stage of growth, under water by means of an arrangement (net?), and feeding them with finely chopped earthworms, he was able to keep them for several months—and indeed throughout the winter—in this condition, and in this way to forcibly defer their final change, and their transition from the tadpole stage to that of the perfected creature during this period.” It is not stated whether the animals finally underwent transformation, so that it cannot be decided whether we have here a case of reversion or simply one of retarded development. That metamorphosis may occur after a long period of time, is shown by experiments upon the tadpole of Pelobates conducted by Professor Langer in Vienna.[264] The creatures were kept in deep water in such a manner that they were not able to land, and by this means three out of a large number of individuals had their metamorphosis delayed till the second summer; notwithstanding this, transformation then occurred.

It cannot be objected to my reversion hypothesis, that it opposes on the one side what on the other it postulates, viz. a per saltum change of structure. Reversion is characterized by the sudden acquisition of an older, i.e. a formerly existing phyletic stage. That reversion occurs is a fact, whilst nobody has hitherto been able to prove, or even to make probable, that a stage of the future (sit venia verbo) has been attained at once (per saltum).

Now if it is possible to find influences in the present conditions of life of the Axolotl which make it difficult or quite impossible for it to live upon land, and which therefore appear as incentives to the reversion to the Ichthyodeous form, the other portion of my hypothesis—the assumption that the Axolotl had become an Amblystoma at a former period—can also be supported by facts.

We know from Humboldt[265] that the level of the Lake of Mexico at a comparatively recent period was considerably higher than at present. We know further that the Mexican plateau was covered with forest, which has now been destroyed wherever there are human, and especially Spanish settlements. Now if we suppose that at some post-glacial period the mountain forests extended to the borders of the lake, at that time deep, with precipitous sides and much less saline, not only should we thus have presented different conditions of life to those at present existing, but also such as would be most favourable for the development of a species of salamander.

On the whole, I believe that my attempt to explain the exceptional metamorphosis of the Axolotl of the Mexican lake cannot be objected to as being a too airy phantasy. In any case it is the only possible explanation which can be opposed to that which supposes that the occasional transformation of the Axolotl is not reversion, but an attempt at advancement. This last assumption must, in my judgment, be rejected on purely theoretical grounds by those who hold that a sudden transformation of a species, when connected with adaptation to new conditions of life, is inconceivable—by those who regard adaptation, not as the sudden work of a magic power, but as the end result of a long series of natural, although minute and imperceptible causes.

If my interpretation of the facts be correct, there arises certain consequences which I may here briefly mention in conclusion.

First, with regard to more obvious results. If Siredon Mexicanus, Shaw, only by occasional reversion assumes the Amblystoma form, and never, or only exceptionally, propagates as such, but only as Siredon, the more recent systematists are not justified in striking out the genus Siredon and in placing S. Mexicanus as an undeveloped form in the genus Amblystoma. So long as there exists not one only, but several species of Siredon which as such regularly propagate themselves, the genus exists; and although we would not deprive systematists of all hope of these species of Siredon being one day re-elevated to Amblystomæ, it nevertheless better accords with the actually existing state of affairs if we allow the genus Siredon to remain as before among the genera of Salamandrina, and to include therein all those species which, like the Paris Axolotl, S. Mexicanus, Shaw, and probably also S. Lichenoides, Baird, only exceptionally, or through artificial influences, assume the Amblystoma form, but without propagating regularly in this condition. On the other hand, we should correctly comprise under the genus Amblystoma all those species which propagate in this state regularly, and in which the perennibranchiate stage occurs only as a larval condition.

To arrive at a decision in single cases would chiefly concern the American naturalists, whose ever increasing activity may lead us to hope soon for a closer investigation of the reproduction of the numerous species of Amblystoma of their native country. I should rejoice if the facts and arguments which I have here offered should give an impetus to such researches.

The second consequence to which I may refer, is of a purely theoretical nature, and concerns a corollary to the “fundamental biogenetic law” first enunciated by Fritz Müller and Haeckel. This, as is well known, consists of the following law:—The ontogeny comprises the phylogeny, more or less compressed and more or less modified.

Now according to this law, each step in phyletic development when replaced by a later one, must remain preserved in the ontogeny, and must therefore appear at the present time as an ontogenetic stage in the development of each individual. But my interpretation of the transformation of the Axolotl appears to stand in contradiction to this, since the Axolotl, which at a former period was an Amblystoma, retains nothing of the latter in its ontogeny. The contradiction is, however, only apparent. As long as we are concerned with an actual advance in development, and therefore with the attainment of a new step never formerly reached, the older stages will be found in the ontogeny. But this is not the case when the new stage is not an actual novelty, but formerly represented the final stage of the individual development; or, in other words, when we are concerned with the reversion, not of single individuals, but of the species as such, to the preceding phyletic stage, i.e. with a phyletic degeneration of the species. In this case the former end-stage of the ontogeny would be simply eliminated, and we should then only be able to recognize its former existence by its occasional appearance in a reversion form. Thus, under certain conditions the Triton sinks back to the perennibranchiate stage; not in such a manner that the individual first becomes a Triton and then undergoes perennibranchiate re-modification, but simply, as I have already shown above, by its remaining at the Ichthyodeous stage and no longer attaining to the Salamander form. So also, according to my hypothesis, the salamandrine Amblystoma Mexicanum, formerly inhabiting the shores of the Lake of Mexico, has degenerated to the perennibranchiate stage, and the only trace that remains to us of its former developmental status is the tendency, more or less retained in each individual, to again ascend to the salamander stage under favourable conditions.

The third and last consequence which my interpretation of the facts entails, is the change in the part played by reversion in organic nature. Whilst atavistic forms have hitherto been known only as isolated and exceptional cases, interesting indeed in the highest degree, but devoid of significance in the course of the development of organic nature, a real importance in this last respect must now be attached to them.

I may assume that reversion can in two ways be effectual for the preservation or re-establishment of a living form. In the first place, where, as in Axolotl, the new and organically higher form becomes untenable through external influences, instead of simply perishing—since advancement in another direction does not appear to be possible—a reversion of the species to the older and more lowly organized stage occurs. In the second place, the older phyletic form may not be abandoned while a newer form is being developed therefrom, but the former may alternate with the latter, as we see in the case of seasonally dimorphic butterflies. It can hardly be objected if I regard the alternation of the summer and winter form in this case as a periodic reversion to the phyletically older (winter) form.

Although the reversion of an entire species, such as I suppose to have been the case with the Axolotl, may be of rare occurrence, this is certainly not the case with periodic or cyclical reversion; the latter plays a very important part in the development of the various forms of alternating or cyclical propagation.[266]

Postscript.

In the previous portion of this essay it was pointed out that the causes to which I attributed the reversion of the hypothetical Amblystoma Mexicanum to the existing Axolotl, did not appear to me to amount to a complete explanation of the phenomenon. In the first place these seemed to me too local, since they could only be applied with any certainty to the Axolotl of the lake of the Mexican capital, whilst the Paris Axolotls obtained from other parts of Mexico still required an explanation. On the other hand, these causes did not appear to me sufficiently cogent. Should we even learn subsequently that the Paris Axolotl is also derived from a salt lake which is exposed to similar winds to the Lake of Mexico, we still have in this peculiarity of the lakes only a cause tending to make it difficult for the larva to undergo metamorphosis, and to reach a suitable new habitat on the land. The impossibility of doing this, or the complete absence of such habitat, does not however follow as a necessary consequence.

It would obviously be a much more solid support for my hypothesis if it were possible to point to some physical conditions of the land which there precluded the possibility of the existence of Amblystomas.

For a long time I was indeed unable to discover such causes, and I therefore concluded the previous portion of this essay and went to press. Afterwards, when residing in one of the highest valleys of our Alps in the Upper Engadine, an idea accidentally occurred to me, which I do not now hesitate to regard as correct after having tested it by known facts.

It happens that in the Upper Engadine there live only such Amphibia as persistently, or at least frequently resort to the water. I found frogs up to nearly 7000 feet above the sea, and Tritons at 6000 feet (Pontresina and Upper Samaden). On the other hand, the land-living mountain salamander, S. Atra,[267] was absent, although suitable stations for this species were everywhere present, and it would have wanted for food as little as do its allies the water-newts. Neither would the great elevation above the sea offer any obstacle to its occurrence, since it occasionally ascends to a height of 3000 metres (Fatiot).[268]

Now it is well known that the atmosphere of the Upper Engadine,[269] like that of other elevated Alpine valleys enclosed by extensive glaciers, is often extraordinarily dry for a long period, a condition which appears to me to explain why the black land-salamander is there absent,[270] whilst its near water-living ally occurs in large numbers. The skin of the naked Amphibia generally requires moisture, or else it dries up, and the creature is deprived of a necessary breathing apparatus, and often dies as rapidly as though some important internal organ had been removed. Decapitated frogs hop about for a long time, but a frog which escapes from a conservatory and wanders about for one night in the dry air of a room, is found the following day with dry and dusty skin half dead in some nook, and perhaps perishes in the course of another day if left without moisture.

All that we know of the biology of the Amphibia is in accordance with this. Thus, all the land-salamanders of southern Italy avoid the hot and dry air of summer by burying in the ground, where they undergo a summer sleep. This is the case with the interesting Salamandrina Perspicillata,[271] and with the land-living Sardinian Triton, the remarkable Euproctus Rusconii, Gené,[272] (Triton Platycephalus, Schreiber). With respect to Geotriton Fuscus I learn from Dr. Wiedersheim, who has studied the life conditions of this, the lowest European Urodelan, in its own habitat, that in Sardinia it sleeps uninterruptedly from June till the winter; whilst on the coast of Spezia and at Carrara, where it also occurs, it avoids the summer sleep in a very peculiar manner. It makes use of the numerous holes in the calcareous formation of that region, and for some months in the year becomes a cave-dweller. As soon as the great heat occurs, often in May, it withdraws into the holes, and again emerges in November during the wet weather. In these lurking holes it does not fall into a sleep, but is found quite active, and its stomach, filled chiefly with scorpions, shows that it goes successfully in search of food; the moist air of the holes makes it unnecessary for it to bury in the earth.

In the same sense it appears to me must be conceived the fact that the solitary species of frog of the Upper Engadine, Rana Temporaria,[273] the brown grass frog, is there much more a frequenter of the water than in the plains. It is true that I can find no remark to this effect in the excellent work of Fatiot, already referred to above, and I am therefore obliged to resort to my own observations, which, although often repeated, have always been carried on for only a short time. I was much struck with the circumstance that the Engadine frogs were to be found in numbers in the water long after the pairing season, which, according to Fatiot, lasts at most to the end of June. In the numerous pools around Samaden I found them in July and August, whilst in the plains they only take to the water at the time of reproduction, and seek winter quarters in the mud on the first arrival of this season. (Fatiot, p. 321.) In the Engadine they have therefore in some measure adopted the mode of life of the aquatic frogs, but this of course does not prevent them from returning in damp weather to their old habits and roving through meadows and woods.

After these considerations had made it appear to me very probable that the dry air of the Upper Engadine accounted for the absence of the black land-salamander, the question at once arose whether the absence of Amblystomas from the Mexican plateau might not perhaps be due to the same cause, i.e. whether such a dryness of the atmosphere might not perhaps prevail also in that region, so that Amphibia, or at least salamander-like Amphibia, could not long exist on the land. The height above the sea is still greater (7000 to 8000 feet), and the tropical sun would more rapidly dessicate everything in a country poor in water.

As I was at the time without any books that might have enlightened me on the meteorological conditions of Mexico, I wrote to Dr. v. Frantzius, who, by many years residence in Central America was familiar with the climate of this region, and solicited his opinion. I received the reply that on the high plains of Mexico an extraordinary dryness of the atmosphere certainly prevails. “The main cause of the dryness of the high plains is to be found in the geographical position, the configuration of the land, and the physical structure. The north-eastern trade-wind drives the clouds against the mountains, on the summits of which they deposit their moisture, so that no vapour is carried over; as long as the north-east trade-wind blows, the streams feeding the rivers flowing into the Atlantic Ocean are abundantly fed with water, whilst on the western slopes, and especially on the high plains, the clouds give no precipitation. In the second half of the year also, during our summer, the so-called rainy season brings but little rain[274]—little in comparison with the more southern regions, where the heavy tropical thunderstorms daily deluge the earth with water. Mexico lies much too northerly, and does not reach the zone of calms, within which region these tropical rains are met with.”

Thus, in the high degree of dryness of the air lasting throughout the year, I do not doubt that we have the chief cause why no Amblystomas occur on these elevated plains; they simply cannot exist, and would become dried up if taken there, supposing them not to be able to change their mode of life and to take to the water. If therefore in former times Amblystomas inhabited Mexico, the coming on of the existing climatic conditions left them only the alternative of becoming extinct, or of again taking to the aquatic life of their Ichthyodeous ancestors. That this was not directly possible—that the Amblystoma form was not able to become aquatic without a change of structure, is shown by the fact that even in the Lake of Mexico no Amblystoma occurs. A retreat to an aqueous existence could, as it appears, only be effected by complete reversion to the Ichthyodeous form, which then also took place.

But my hypothesis of the transformation of the Axolotl not only requires the proof that Amblystomas cannot exist under present conditions in Mexico, but also the further demonstration that at a former period other conditions prevailed there, and these of such a nature as to make the existence of land-salamanders possible.

With respect to my question, whether we might not perhaps assume that at some post-glacial period the conditions of atmospheric moisture on the high plains of Mexico were essentially different from those at present prevailing, I recollected Dr. v. Frantzius and the above-quoted observation of Humboldt’s,[275] who discovered in the neighbourhood of the Lake of Tezenco (Mexico) distinct evidence of a much higher former level of the water. “All such elevated plains were certainly at a former period so many extensive water-basins, which gradually became filled, and are still filling up with detritus. The evaporation from such large surfaces of water must at that time have caused a very moist atmosphere, favourable to vegetation and adapted for the life of naked Amphibia.”

From this side also my hypothesis thus receives support, and we may assume with some certainty that at the beginning of the diluvial period[276] the woods surrounding the Mexican lakes were inhabited by Amblystomas, which, as the lakes subsequently became more and more dried up and the air continually lost moisture, found it more difficult to exist on the land. They would at length have completely died out, had they not again become aquatic by reversion to the Ichthyodeous form. It may perhaps be supposed that the above-mentioned physical conditions—desolate, salt-incrusted shores—co-operated in the production of the reversion, by making it difficult for the larvæ to quit the water; but we can only judge with certainty upon this point when, by means of experiment, we have discovered the causes which produce reversion in the Amphibia.

Addendum.

I have lately met with another interesting notice on the reproduction of the native North American Amblystomas. Professor Spence F. Baird, of Washington, has often observed the development from the egg of various species, and especially of Amblystoma Punctatum and A. Fasciatum. His observations do not appear to be as yet published, so that I was unable to discover any account of the development of Amblystoma in existing literature.[277] I am authorized to extract the following brief data from a letter addressed to Dr. v. Frantzius.

In order to deposit their eggs the Amblystomas go into the water, where the eggs are laid enclosed in a jelly-like mass, but never more than fifteen to twenty together. The spherical eggs are very large, perhaps a quarter of an inch in diameter. They soon develop into a Siredon-like larva, which remains several months in this condition. The gills then shrivel up, the creature begins to crawl, and gradually passes through the different transformations to the complete Amblystoma form.

It appears from this communication that the Amblystomas lay much larger and much fewer eggs than the Axolotl, and that their development throughout resembles that of our salamanders.

In concluding I may mention an anatomical fact which most strongly supports my view that the Mexican Axolotl is a reverted Amblystoma. I learn from Dr. Wiedersheim that the Axolotl possesses the “intermaxillary gland” which occurs in all the land Amphibia. This organ, lying in the intermaxillary cavity, appears, whenever it occurs, to produce a kind of birdlime, i.e. a very glutinous secretion, which serves to attach the prey to the rapidly protrusible tongue. Although this secretion may perhaps also have another function, from the absence of the intermaxillary gland in all exclusively aquatic Amphibia, it follows that it must be devoid of importance for, and inapplicable to feeding in the water. The intermaxillary gland is absent in all Perennibranchiata and Derotremata which Wiedersheim has hitherto investigated, viz. in Menobranchus, Proteus, Siren, Cryptobranchus, Amphiuma, and Menopoma, all of which are indeed without the cavity in which the gland is situated in the Salamandrina, i.e. the cavum intermaxillare.

Now in the Salamandrina the gland appears at an early stage. It is possessed in a well-developed state by the larvæ both of species of Triton and of Amblystoma, where indeed the glandular structure completely fills the cavum intermaxillare.

Were the Axolotl a species retarded in phyletic development, the presence of a gland which does not occur in any other Perennibranchiata, and which is only of use for life upon land, would be quite inexplicable.

The matter becomes still more enigmatical through the fact that the gland, although present, is quite rudimentary. Whilst in the Salamandrina the capacious intermaxillary cavity is entirely filled by the tubes of the gland in question, in Axolotl this cavity is almost completely filled with a closely woven connective tissue, in which there can only be found a small number of gland-tubes—in the extreme front, and at the base immediately over the intermaxillary teeth—these tubes agreeing in the details of their histological structure with the elements of the same gland in the Salamandridæ.

I give these anatomical details from Dr. Wiedersheim’s verbal communication. An amplified account will subsequently appear in another place.[278]

An explanation of this rudimentary intermaxillary gland in the Axolotl only appears to me possible on the supposition that the latter is an atavistic form. From this point of view it is evident that the gland already present in all Amblystoma-larvæ must have been taken over by the perennibranchiate form of the existing Axolotl, through the reversion of the hypothetical Amblystoma Mexicanum of the “diluvial period.”[279] It can also be easily understood that this organ would become more and more rudimentary in the course of time, since it has no further use in the water, and the gap thus arising in the formerly present cavum intermaxillare would become filled with connective tissue.

While the German edition of this work was going through the press I obtained, through the kindness of my friend Dr. Emil Bessels of Washington, the Mexican memoir upon the new Axolotl,[280] which even in Mexico regularly, or at least in many cases, becomes developed into the Amblystoma form.

The facts are briefly as follows:—The small Lake of Santa Isabel is some hours’ journey from the Mexican capital. In this lake there lives a species of Axolotl which had hitherto remained unknown, and was described by Señor Velasco as Siredon Tigrinus. This species propagates itself indeed in the Axolotl state, but in many cases it becomes transformed into Amblystoma and takes to the land. Although propagation in the Amblystoma condition was not observed, it can hardly be doubted that it also propagates in this form.

At first sight these facts appear to refute my hypothesis, that the extreme dryness of the air of the Mexican plateau precludes the existence of land Amphibia. Nevertheless I do not abandon this hypothesis for the former one, since a closer study of the data furnished by Velasco confirms rather than refutes my supposition.

Velasco expressly corroborates the statement that the Axolotl hitherto known from the great Mexican lake which never dries up (Lake of Xochimilco and Chalco), is only met with in its native habitat in the Siredon form, i.e. as Siredon Humboldtii. According to Velasco the cause of the frequent assumption of the Amblystoma form by the new Siredon Tigrinus, is to be found in the local conditions of life of this species. The Lake Santa Isabel is shallow, its greatest depth amounting to three meters, and it is liable to a periodical drying up, which is so complete that one can pass dry-shod through it in several places. The species must therefore have long since died out had it not been able to adapt itself periodically to a land life. Now it could have become transformed into a land Amphibian—as Señor Velasco observed—at various stages of growth; and indeed this author believes that “the Creator has implanted an instinct in this creature,” which enables it to always undergo metamorphosis at the right time.

This last assumption may or may not be taken as correct, but this much is established, viz. that numerous individuals of this species take to the land, and remain there during a period of many months.

But does this contain the proof that salamander-like animals are actually able to lead a land life in Mexico—that the dry air is advantageous, or at least supportable to them? It does not appear so to me, but rather that all which has been reported of this Amblystoma by Señor Velasco goes to show that the animal does not, properly speaking, live upon land like the North American Amblystomas, or like our land-salamanders, but that it only experiences a summer sleep lasting over the period of drought. These Amblystomas were observed as they left the dried-up lake at night in order to seek some moist lurking-place in the neighbourhood, where they might remain concealed. They are only known in the villages situated near the lake, and were only seen there at large just when they were wandering from the lake to their place of concealment. At other times they were mostly found in the earth, buried under walls, the pavement of the market-place, &c. When laying down a line of railway, a workman found in the earth a whole nest of twelve Amblystomas lying close together. All these are not mere lurking-holes which could be abandoned at any moment; it would rather appear that we have here places of refuge for the entire duration of the period of drought, and that these would only be forsaken when the water of the rainy season penetrated the soil. I am not myself in a favourable position for investigating these suppositions more closely, but this could be done by Señor Velasco, who lives in Mexico, and science would be much indebted to him if he would examine as precisely as possible into the habits and conditions of life of this, and of the other species of Mexican Axolotls. Unfortunately this gentleman can, it would appear, have seen only the French publications upon the transformation of the Axolotl, and could not therefore have asked himself questions arising from my conception of the facts; otherwise many of his observations would have led to more definite results. The above conclusion can however be still further supported by Señor Velasco’s data.

One might indeed insist that with us also the land-salamanders conceal themselves in moist places during dry weather, and often lie hidden, as in Mexico, in a hole, in a cluster of as many as ten together; but with us they leave their lurking-place from time to time and go in search of food. Señor Velasco mentions nothing with respect to this. What especially struck me was the statement that the Mexican Amblystomas were also to be found in the water.[281] When Lake Santa Isabel is drained, the fishermen stretch large nets across the exit channels, and in these they not only find ordinary Axolotls, but also some “sin aretes,” which they also designate “mochos,” i.e. hornless Axolotls, because they have no gills, but have already reached the Amblystoma stage. Our land-salamanders live in the water only as larvæ, but they also love and require moisture. Only the female enters the water when she wants to deposit her young (eggs with mature larvæ), and then only at the margin of shallow pools or small brooks. The Mexican Amblystoma thus much more resembles in its habits our water-salamanders (Tritons), which remain in the water at least during the whole period of reproduction. These also leave the water later, and, like the land-salamander, seek concealment in the earth. They have this habit also in those districts which possess a very dry atmosphere; and especially in the Engadine, where I first conceived the idea of taking into account the dryness of the air, I found in the pools at the end of August and the beginning of September only larvæ of Tritons. The older Amphibians must therefore have been on the land, presumably in their places of winter concealment.

From what we have hitherto learnt from Señor Velasco, the mode of life of Amblystoma Tigrinum must resemble that of our Tritons, although its structure is that of a land-salamander. I would thus offer the following explanation of the facts at present known:—Owing to the periodic drying up of the lake of Santa Isabel, the Siredon Tigrinus would be again compelled to undergo metamorphosis. Whether this was formerly entirely abandoned, or whether it always occurred in solitary individuals, is almost immaterial; in any case the habit of metamorphosis must have been very rapidly acquired through natural selection, and must have again become general, if the faculty was only present in the species, although latent. Through the dryness of the air, the Amblystomas that had taken to the land would be compelled to bury themselves at once, and to remain asleep till the recurrence of the rainy season, when they would hasten back into the water and would there live as a species of Triton.

Now one might feel inclined to ask why the species of the great Mexican lake has not also taken to this mode of life. To this it may be simply replied that the water of this lake never dries up, and that the Axolotls have thus never been reduced to the alternative of undergoing metamorphosis or of perishing. If therefore the conditions of existence in water were more favourable than on land, the tendency to abandon metamorphosis would increase from generation to generation, and the deportment at present observed would finally result, i.e. propagation would take place exclusively in the Axolotl state. As has already been mentioned above, the latest observations of Velasco furnish further confirmation that the Axolotl of the great lake is never met with in the Amblystoma condition, “although it (the Axolotl) is brought daily from Mexico into the market throughout the whole year.” I should not however regard it as a refutation of my view if prolonged investigation should show that this species also (Siredon Humboldtii) occasionally developed into an Amblystoma; on the contrary, it would not at all surprise me if such cases of reversion occurred in Mexico as well as in Europe. The fact that an immense majority of the Amphibians propagate in the Axolotl state would not be thereby affected, and would still require an explanation: this I am still inclined to see in the dryness of the air of the high plains, which is so unfavourably adapted for a life passed entirely on land.

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IV.
ON THE MECHANICAL CONCEPTION OF NATURE.

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INTRODUCTION.

In the first of the three preceding essays it was attempted to solve the question whether the transformations of a given complex of characters in a certain systematic group could be completely explained by the sole aid of Darwinian principles. It was attempted to trace the origin of the marking and colouring of the Sphinx-caterpillars to individual variability, to the influences of the environment, and to the laws of correlation acting within the organism. These principles as applied to the origin of a certain well-defined, although narrowly restricted range of forms, were tested in order to see whether they were alone sufficient to explain the transformation of the forms.

It appeared that this was certainly the case. In all instances, or at least where the facts necessary to obtain a complete insight were available, the transformations could be traced to these known factors; there remained no inexplicable residual phenomena, and we therefore had no reason for inferring the existence of some still unknown modifying cause lying concealed in the organism. In this region of the marking and colouring of caterpillars, the assumption of a phyletic vital force had to be abandoned, as being superfluous for the explanation of the facts.

In the second essay the attempt was next made with reference to double form-relationship, as presented for observation in metamorphic insects, to draw conclusions as to the causes of the transformations. It appeared here that form- and blood-relationship do not always coincide, since the larvæ of a species, genus, or family, &c., may show quite different form-relationships to their imagines. These facts alone told very decisively against the existence of an internal developmental power, so that the latter had likewise to be set aside by the method of elimination, since the observed incongruences as well as the congruences of form-relationship, found sufficient explanation in the action of the environment on the organism.

This investigation thus also led to the denial of a phyletic vital force.

In the third essay I finally sought to prove that the only case of transformation of one species into another at present actually observed[282], could not without further evidence be interpreted as the result of the action of a phyletic vital force, but that more probably we had here only an apparent case of new formation, which was in reality but a reversion to a stage formerly in existence.

If this last investigation removes the only certain observation which could have been adduced in favour of the hypothesis of a phyletic vital force, so also do the two former essays show that this hypothesis, at least in the case of insects, must be abandoned as inadequate.

The question now arises whether this conclusion, based on such a limited range of inquiry, can also be applied to the other groups of the organic world without further evidence.

The supporters of a principle of organic development will deny this in each individual case, and will demand special proof for each group of organisms; I believe this position, however, to be incorrect. Here, if anywhere, it appears to me justifiable to apply the conclusions inductively from special cases to general ones, since I cannot at all see why a power of such pre-eminent and fundamental importance as a phyletic vital force should have its activity limited to solitary groups in the organic world. If such a power exists it must be the inciting cause of organic development in general, and must be equally necessary in every part of creation, as no advancement could take place without it. In this case, however, the force would be recognizable and demonstrable at every point; the phenomena should nowhere stand in opposition to its admission, and should in no case be explicable or comprehensible without it. The same laws and forces which caused the development of one group of forms must underlie the development of the whole organic world.

I therefore believe that we are correct in applying to the whole living world the results furnished by the investigation of insects, and in thus denying the existence of an innate metaphysical developmental force.

There is, however, a quite distinct method which leads to the same results, and to the preliminary, if not to the complete and definitive rejection of such a principle; the admission of this power is directly opposed to the laws of natural science, which forbid the assumption of unknown forces as long as it is not demonstrated that known forces are insufficient for the explanation of the phenomena. Now nobody will assert that this has in any case been proved; the test of applying the known factors of transformation has only just commenced, and wherever it has been made they have proved sufficient as causal forces. Thus, even without the foregoing special investigations we should deny a phyletic vital force; the more so as its admission is fraught with the greatest consequences, since it involves a renunciation of the possibility of comprehending the organic world. We should, on this assumption, at once cut ourselves off from all possible mechanical explanation of organic nature, i.e. from all explanation conformable to law. But this signifies no less than the renunciation of all further inquiry; for what is investigation in natural science but the attempt to indicate the mechanism through which the phenomena of the world are brought about? Where this mechanism ceases science is no longer possible, and transcendental philosophy alone has a voice.

This conception represents very precisely the well-known decision of Kant:—“Since we cannot in any case know à priori to what extent the mechanism of Nature serves as a means to every final purpose in the latter, or how far the mechanical explanation possible to us reaches,” natural science must everywhere press the attempt at mechanical explanation as far as possible. This obligation of natural science will be conceded even by those who lay great stress upon the necessity for assuming a designing principle. Thus, Karl Ernst von Baer states that we have no right “to assert of the individual processes of Nature, even when these evidently lead to a definite result, that some Mind has originated them designedly. The naturalist must always commence with details, and may then afterwards ask whether the totality of details leads him to a general and final basis of intentional design.”[283]

But even if we are precluded on these grounds only from assuming the existence of a directive power, i.e. a phyletic vital force, for explaining detailed phenomena, and are at the same time debarred from the possibility of arriving at a physical or mechanical explanation—which amounts to no less than the abandoning of the scientific position—it certainly cannot be asserted that the development of the organic world is already conceived of as a mechanical process. We rather acquiesce in the belief that the processes both of organic and of inorganic nature depend most probably upon purely causal powers, and that the attempt to refer these to mechanical principles should not therefore be abandoned. There is no ground for renouncing the possibility of a mechanical explanation, and the naturalist must not therefore resign this possibility; for this reason he cannot be permitted to assume a phyletic power so long as it is not demonstrated that the phenomena can never be understood without such an assumption.

It cannot be raised as an objection that even for the explanation of individual life a vital power was long ago admitted, as there was not then sufficient material at hand to enable the phenomena of life to be traced to physical forces. It is now no longer questionable that this assumption was a useless error—a false method—at the time when made certainly very excusable, since the aspect of the question was then, owing to the imperfect basis of facts, very different to the present analogous question as to the causes of derivative development. Thus, although it is now easy to prove this assumption to be erroneous, it was in the former sense correct, as it was in accordance with the existing state of knowledge. At that time there was hardly one of the numerous bridges which now connect inorganic with organic nature, so that the supposition that life depended upon forces which had no existence outside living beings was sufficiently near.

In any case the philosophers of that period cannot be blamed for filling up the gaps in the existing knowledge by unknown powers, and in this manner seeking to establish a finished system. The task of philosophy is different to that of natural science; the former strives at every period to set up a completely finished representation of the universe in accordance with the existing state of knowledge. Natural science on the other hand is only concerned in collecting this knowledge; she need not therefore always finish off, and indeed can never close her account, since she will never be in a position to solve all problems.[284] But science must not for this reason pronounce any question to be insoluble simply because it has not yet been completely solved; this she does, however, as soon as she renounces the possibility of a mechanical explanation by invoking the aid of a metaphysical principle.

That this is the correct mode of scientific investigation is seen by the abandoning of the (ontogenetic) vital force. The latter is no longer admitted by anybody, now that we have turned from mere speculation to the investigation of Nature’s processes; nevertheless its non-existence has not been demonstrated, nor are we yet in a position to prove that all the phenomena of life must be traced to purely physico-chemical processes, to say nothing of our being actually able to thus trace them. Von Baer also states “that the abolishment of the vital force is an important advance; it is the reduction of the phenomena of life to physico-chemical processes, although these indeed still contain many gaps.” He points out how very far we are still removed from being able to reduce to physical causes, the processes through which the fertilized yelk of an egg becomes developed into a chicken.

How comes it therefore that we all have a conviction that such a complete reduction will in time become possible, or if not this, that the development of the individual depends entirely upon the same forces which are in operation without the organism? For what reason have we rejected the “vital force”?

Simply because we see no reason for assuming that known forces are insufficient for explaining the phenomena, and because we are not justified in admitting directive forces as long as we have any hope of one day furnishing a mechanical explanation.

But if it is not only permissible, but even necessary, to explain the ontogenetic vital power by known forces, and to commence to indicate the mechanism which produces the individual life, why should it not be equally necessary to abandon that assumption of a phyletic vital force which stifles any deeper inquiry, and to attempt to point out that here also the co-operation of mechanical forces has brought about the multitudinous and wonderful phenomena of the organic world?

The renunciation of the old vital force was certainly an immediate consequence of the acquisition of new facts—of the knowledge that the same compounds which compose organic bodies can be produced without the latter. This discovery, due to Wöhler and his followers, showed that organic products could be prepared artificially.[285] In brief, the decline of the vital force followed from the knowledge that at least one portion of the processes of life was governed by known forces.

But in the domain of the development of the organic world have we not quite analogous proofs of the efficacy of known forces? Is not the variability of all types of forms a fact? and must not this under the action of natural selection and heredity lead to permanent changes? Has not the problem of explaining the subserviency of all organic form to law as a result without invoking its aid as a principle been thus successfully solved? It is true that we have not directly observed the process of natural selection from beginning to end; neither has anybody directly observed the mode in which the heat of the animal body is generated by the processes of combustion going on in the blood and in the tissues; nevertheless, this is believed as a certainty, and a “vital force” is not invoked.

Now the above-mentioned Darwinian principles of transmutation are certainly not simple forces of nature like those underlying the development of the individual, i.e. chemico-physical forces, and it cannot be said à priori whether in one of these principles—perhaps in variability or in correlation—there may not lie concealed a metaphysical principle in addition to the physical forces. In fact it has lately been asserted by Edward von Hartmann[286] that the theory of selection is not a mechanical explanation, since it combines forces which are only partly mechanical and in part directive.

It must therefore be next investigated whether this assertion is tenable.

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I.
Are the Principles of the Selection Theory Mechanical?

Edward von Hartmann may justly claim that his views should be considered and tested by naturalists.[287] He would be correctly classed with those philosophers who have approached this question with a many-sided scientific preparation. It can nevertheless be perceived in his case how difficult, and indeed how impossible, it is to estimate the true value of the facts furnished by the investigation of nature, when we attempt to take up only the results themselves, without being practised in the methods by which these are reached, i.e. without being completely at home in one of the scientific subjects concerned through one’s own investigations. It appears to me that the denial of the purely mechanical value of the Darwinian factors of transformation arises in most part from an erroneous classification of the scientific facts with which we have to deal. There can certainly be no mistake that the entire philosophical conception of the universe, as laid down by Von Hartmann in his “Philosophy of the Unconscious,” is unfavourable to an unprejudiced estimate of scientific facts and to their mechanical valuation.

Variability, heredity, and above all correlation, would not be regarded by Von Hartmann as purely mechanical principles, but he would therein assume a metaphysical directive principle.

In the first place, as regards variability, Von Hartmann endeavours to show that it is only a quite unlimited variability which suffices for the explanation of necessary and useful adaptations by means of selection and the struggle for existence. But this does not exist—variation rather takes place in a fixed direction only (in Askenasy’s sense), and this can be nothing else than the expression of an innate law of development, i.e. a phyletic vital force.

This deduction appears to me in two ways erroneous. In the first place it is incorrect that a quite unlimited variability is a postulate of the theory of selection, and in the next place the admission of variability, which is in a certain sense “fixed in direction,” does not necessitate the assumption of a phyletic vital force.

A mere unsettled variability, uniform in all possible directions, is, according to Von Hartmann, necessary for the theory of selection, because only then does the variability offer a certain guarantee “that under given conditions of life the variations necessary for complete adaptation will not be wanting.” But it is hereby overlooked that the new life conditions to which the adaptation must take place are as little fixed and unchangeable as the organism itself. In such a case of transformation we have not to deal with a type of organization which was before fixed and immutable, and which has to be squeezed into new life-conditions as into a mould. The adaptation is not one-sided, but mutual; a species in some measure selects its new conditions of life, corresponding with those possible to its organization, i.e. with the variations actually occurring. I will choose an instance which will even be conceded by Von Hartmann as being only explicable by natural selection, viz., a case of mimicry.

Supposing that among the South American Heliconiidæ there occurred a species of Pieris which had no resemblance to these protected butterflies, either in form, marking, or colouring; who can deny that it would be most useful to this species to acquire the form and colouring of a Heliconide, and thus, by taking to new conditions of life, to avoid the persecutions of its foes? But if the physical nature of the Pieride concerned precluded the occurrence of Heliconoid variations, would this incapability of insinuating itself into these new conditions necessitate the decline of the species? Could not its existence be secured in some other manner? could not the destruction of numerous individuals by foes be compensated for by increased fertility? to say nothing of the numerous other means through which the number of surviving individuals might become increased, and the existence of the species secured. This case is not arbitrarily chosen; in the districts where the Heliconiidæ occur there are actually a large number of Whites which do not possess the protective colours of the former nauseous family. In the adoption of these new life conditions we have not to deal therefore with survival or extermination, but only with amelioration. It is not every species of “White” that can become adapted to these conditions, because every species does not give rise to the necessary colour variations; those that do, become in this way modified, because they are thus better protected than before. And so it is throughout; wherever we find protected insects enjoying immunity from foes we see also mimickers, sometimes only single, sometimes several, and generally from very diverse groups of insects, according to the general resemblance which existed before the commencement of the process of adaptation, and to the variations made possible by the physical nature of the species concerned.

In the first essay of the second part of this work it was shown that in certain Lepidopterous larvæ a process of adaptation is at the present time still in progress, this depending upon the fact that while the young caterpillar is very well protected by the leaf-green colour of its body, this colour becomes insufficient to conceal the insect as soon as it exceeds the leaf in size. All such caterpillars—and there is a whole series of species—as they increase in size acquire the habit of concealing themselves on the earth by day, and of feeding only at night. New conditions of life are thus imposed, and these are even compulsory, i.e. they could not be abandoned without risking the existence of the species. Now in accordance with these new conditions, some individuals in these species have lost the green colouring of the young stages, and have acquired the brown coloration of the dark surroundings of the insects which conceal themselves by day. In one species this change has now occurred in almost all individuals, in others in only a larger or smaller proportion of them. Now supposing that among these species there occurred one, the physical nature of which did not admit of the production of brown shades of colour, would the species for this reason succumb? Is it not conceivable that the want of colour adaptation might be compensated for by better concealment, i.e. by burrowing into the earth, or by a greater fertility of the species, or by the development of warning signals—supposing the species to be unpalatable—or finally, by the acquisition of a terrifying marking? In other words, could not the caterpillar itself modify the new condition of life—that of being concealed by day—in accordance with variations made possible by its physical nature?

As a matter of fact in one of these species the green colour remains unchanged in spite of the altered mode of life, and this species, wherever it occurs, notwithstanding the persecution of entomologists, is always common (Deilephila Hippophaës); it conceals itself better and deeper however than those other species which, like Sphinx Convolvuli, are difficult to detect on account of their brown colour. In another species the striking yellowish green colouring is likewise retained in the majority of individuals, but this species buries itself by day in the loose soil (Acherontia Atropos).

To this it may be objected that there are also compulsory changes in the conditions of life from which the species cannot withdraw itself, but in which adaptation must necessarily follow, or extermination would take place.

Such compulsory conditions of life do most assuredly occur, and there is indeed no doubt that many living forms have perished through not becoming transformed. I believe, however, that such conditions occur much more rarely than one is inclined to admit at first sight. As a rule the alternative of immediate change or of extermination is offered only by such changes in the conditions of life as occur very rapidly. The sudden appearance of a new and dominant enemy, such as man, has already caused the extinction of the Dodo (Didus ineptus), and of Steller’s Sea Cow (Rhytina Stelleri), and of other vertebrate animals, and constantly leads to the extermination of many other species of different classes. When in America hundreds of thousands of acres of primeval forest are annually destroyed, the conditions of life of a numerous fauna and flora must be thereby suddenly changed, leaving no choice but extermination.

Such abrupt changes in the conditions of life occur, however, but seldom in nature unless caused by man, and must therefore have very rarely happened in former epochs of the earth’s history. Even climatic changes, which we might at first regard as of this character, and which produce a modification in one fixed direction, occur always so gradually that the species has time either to adapt itself to the conditions in this or that direction, according to the variations possible to its physical nature, or else to emigrate.

It thus appears to me erroneous to suppose that variability must be “merely undetermined” in order to complete its part in Darwin’s theory of selection, and its “illimitedness” seems to me also as little necessary for this purpose. Von Hartmann imagines that it is only unlimited variability that furnishes a guarantee that any type, to whatever extent diverging from its point of departure, will be reached by the Darwinian method of gradual transmutation by means of selection and the struggle for existence.

But who has ever asserted that any type can be reached from any point? Or if anybody has said such nonsense, who can prove that its admission is necessary for the theory of selection? Nowhere in systemy do we see any point of support for such an assumption. But when Von Hartmann imagines that the “unlimited” variability which he postulates for Darwin “is in itself unlimited, the limits of its divergence in a given direction being found, not in itself, but only in external obstacles,” he conceives variability to be something independent of, and in some way added to, the animal body, and not a mere expression for the fluctuations in the type of the organism. If, however, we conceive variability in this latter, the true scientific sense, it is in no way “quantitatively unlimited,” nor are its limits even determined by external influences, but essentially by internal influences, i.e. by the underlying physical nature of the organism. Darwin has indeed already shown this in a most beautiful manner in his investigations upon the correlations of organs and systems of organs of the body. To make use of a metaphor, the forces acting within the body are in equilibrium; if one organ becomes changed this causes a disturbance in the forces, and the equilibrium must be restored by changes in other parts, and these again entail other modifications, and so forth. Herein lies the reason why the primary change cannot exceed a certain amount if the restoration of the equilibrium is not to be quite impossible. This is but a metaphor, and I do not wish to assert that we are at present in a position to formulate and demonstrate mathematically for any particular case, how much an organ can become changed in any one species before an interruption of the internal harmony of the body takes place. But such impossibility of demonstration does not appear to me to furnish a sufficient reason for regarding variability as the expression of a directive power—as an “innate tendency to variation conformable to law.”[288] On the contrary, it is to me easily conceivable that we only learn to analyse the processes of nature in detail very slowly, because of their necessary complexity. It thus appears to me quite useless when in this sense Wigand makes use of the objection, that “the gooseberry has not undergone any enlargement since 1852, although it is inconceivable why it should not attain the size of a pumpkin if variability was not internally limited.” It may well be that this is for the present “inconceivable;” nevertheless, this does not justify us in setting up a hypothetical “force of variation” which will not admit of the gooseberry surpassing the pumpkin in size. We are bound to maintain that it is the action and reaction of known forces which sets a limit to the enlargement of this fruit.

In more simple instances the causes of such limitations to growth can be well perceived. Several decades have passed since Leuckart proved in how exact a relation the proportion of volume and surface stood to the degree of organization of an animal. In animals of a spherical form the surface is quite sufficient for respiration, so long as they are of microscopic size. But such an organism cannot become enlarged at pleasure, because the ratio of the surface to the volume would become quite different. The surface increases as the square, whilst the volume increases as the cube, so that very soon the surface of the more rapidly increasing bodily mass can no longer suffice for respiration.[289] This sort of limitation is in no way equivalent to that purely external kind which, for instance, manifests itself in such a manner as to prevent the indefinite lengthening of the tail feathers of the Bird of Paradise. In this case feathers that were too long would hinder flight, and such individuals would accordingly be eliminated by natural selection. The cause is in the former case purely internal, depending upon the equilibrium of the forces governing the organism.

Von Hartmann is entirely in the right when he asserts that variability is neither qualitatively nor quantitatively unlimited. In both senses it is limited (in direction as well as in amount) by the physico-chemical forces acting in some contrary way in each specific organism—by the physical nature of each living form. He errs, however, both in making absolute illimitability a necessary postulate of the theory of selection, as also in inferring the existence of a directive principle from that limitation of variability which is certainly present. “Tendencies to variation” do however exist, not in the sense of a directive power, but as expressions of the different physical constitutions of species, which necessarily cause unequal reactions to the same external actions, as will be more clearly proved below.[290]

This is, of course, a modification of Darwin’s original assumption of an unbounded variability not limited in direction; but Darwin himself has later coincided in the view that the quality of the variations is essentially determined by the nature of the organism.[291]

I now turn to the consideration of the second factor of the theory of selection—heredity. This also, according to Von Hartmann is not a mechanical principle. Darwin himself has now become convinced how great is the probability against the hereditary retention of modifications which, whether feebly or strongly pronounced, appear only in single individuals, i.e. of those so-called “fortuitous” variations which are not the expression of a directive developmental principle. “But as among the numberless possible directions of an indefinite variability, useful modifications can only occur in single cases, Darwin has by this supplementary admission himself retracted an inadmissible assumption of his theory of selection,” and so forth. A “regular, designed tendency to variation, acting from within and contemporaneously affecting a large number of individuals,” must therefore be assumed “in order to insure the by itself improbable inheritance.”

But even from the unbounded variability laid down by the author, it by no means follows that useful variations can only occur in single individuals. In the whole category of quantitative variations the reverse is always the case. Is it the lengthening of some part that is concerned; so would a large number of individuals always possess the useful variation, since we are not dealing with an absolute enlargement, but only with the fact that the part concerned is longer than in other individuals.[292]

But if qualitative variations come into consideration, it may be asked whether Darwin’s “supplementary admission” does not go too far. Such calculations as those quoted by Darwin from the article in the North British Review of March 1867 are extremely deceptive, since we have no means of measuring the amount of protection afforded by a useful variation, and we can therefore hardly compute with any certainty, in how great a percentage of individuals a change must contemporaneously occur in order to have a chance of becoming transferred to the following generation. If our blue rock-pigeon could exist in a polar climate, and if we had the power of introducing it gradually, but not suddenly, into these regions in a wild state, who can doubt that it would assume the white colour of all polar animals? Nevertheless, among wild rock-pigeons white varieties do not occur more frequently than among swallows, crows, or magpies. Or must the white colour of polar animals, the yellow colour of desert species, and the green colour of leaf-frequenting forms, be always referred to a “regular, designed, fixed tendency to variation acting from within,” and causing a “large number of individuals” to vary in a similar manner?

There is, however, a grain of truth in the foregoing; variations which occur singly have but little chance of becoming predominant characters, and this is obviously what Darwin concedes. But this is by no means equivalent to the assumption that only those variations which from the first occur in numerous individuals have a chance of being perpetuated. Let us keep to the facts. We have not the slightest reason either for regarding the white colour of polar animals as the direct action of cold, or for considering that the green colour of foliage-living caterpillars depends upon direct action arising from the habit of resting upon the leaves;[293] both these characters are explicable only by natural selection, and there is nothing to favour the assumption (which Von Hartmann postulates as necessary for success) that many individuals varied into white at the same time. We know no single extra-polar species of a dark colour which frequently, i.e. in many individuals of every generation, varies into white, but we know many species which from time to time produce single white individuals. Now when, on the other hand, we find that all polar animals to which the white coloration is advantageous, and indeed none but species of which the nearest allies vary only individually into white, possess this colour, must we not conclude from this alone that single variations can, under favourable conditions, become predominant characters?

It appears to me that in this question one weighty factor has been too little regarded, even by the supporters of the selection theory, viz., the slowness of most, and especially of climatic changes, which I have already insisted upon. If the transformation of a temperate into an arctic climate occurred so rapidly that the species exposed to it had the alternative either of becoming white in ten or twenty generations or of being unable to exist, then the hasty intervention of a directive power could alone save them from extermination by causing hundreds of thousands of individuals to become similarly coloured with all speed. But it is quite different if the change of climate takes place only in the course of several thousand generations; and this, according to the geological evidence, must have been the true state of the case.

Let us take a definite example—the well-known one of the hare. With us this animal remains brown in the winter and but seldom produces white varieties, whilst its ally the Alpine hare is white during seven months of the year, the Norwegian hare during nine months, and the Greenland hare throughout the whole year. If our climate became transformed into an arctic one, after a given time there would arrive a period when the older coloration no longer possessed any advantage over the occasional and singly-appearing white variations; the winter days during which the ground was covered with snow would have become so numerous, that the protection afforded to the white animals would be equal to the protection enjoyed by the brown individuals on the equally numerous days free from snow. From this time forth the hares that were white in winter would not be subjected to a greater decimation by foxes, &c., than the brown individuals. This period must however be represented as consisting of one or more centuries, and it would be strange if from the individual white hares, which now had an equal chance of existing, some white families did not become established. But the state of affairs would gradually become reversed—the brown hares would experience greater decimation, and wherever there were white families these would possess an advantage in the struggle for existence. It does not follow that the dark individuals would be forthwith extirpated; on the contrary, the advantage in favour of the white would be but small throughout a long period of time, and these individuals would only gradually increase to a higher percentage of the total population; nevertheless their numbers would constantly but very slowly augment. In the course of time this increase would become more rapid for two reasons—first, because even a very small advantage in favour of the increasing number of individuals would always leave a greater number of these victorious; and secondly, because on the whole as the climate became more arctic, the advantage of being white would continually become more decisive in determining which should live and which should succumb.

Thus I see no reason why individual variations which do not appear only once, but which frequently recur in the course of generations, should not acquire predominance under favourable conditions. All facts are in accord with this. Even the common hare shows us that it would be quite capable of becoming coloured in a similar manner. In the museum of Stuttgart there are three specimens of Lepus timidus, killed in Wurtemburg, which are completely white, and several others which are silver-grey or spotted with white. In eastern Russia the common hare possesses a light grey, almost white, winter coat, and Seidlitz[294] makes known the interesting observation that such light specimens occur singly in Livonia, where “the common hare has become naturalized since the commencement of the century.”

As I have already insisted upon above, from the point of view of the conditions of life there is no reason for assuming rapid transformations; the change of conditions is almost always extremely slow; and indeed in numerous instances no objective change occurs, but simply a subjective one, if we may thus designate those cases in which the alteration in the conditions of life depends upon a change in the animal form which is undergoing transformation, and not in that of the environment. This is the case in the above-mentioned instances of mimicry, where the whole change in the conditions of life arises from one species becoming similar to another. The process of natural selection has here as long a period of time as it requires to perfect its results. It is quite similar in all cases of special protective adaptations of form and colour. In all these it is always improvement that is concerned, and not the question “to be or not to be” with which we have to deal.

It is just cases of this last kind, however, which are best fitted for exposing the improbability and insufficiency of the assumption of a variational tendency as a distinct directive power. We have only to fix our attention upon some particular case of sympathetic colouring, or, still better, of mimicry. A “tendency to variation” implies that a large number of individuals produce varieties resembling the model to be imitated, and this—at least according to Von Hartmann—must take place in each of the successive generations, so that by this means, combined with heredity, the useful variation becomes increased. But how comes it that this “tendency to variation” coincides with the existence of the model both in time and place? Can this be due to accident if the two have not a common cause? The upholders of a directive power will certainly not admit this; so that there remains only Leibnitz’s assumption of a pre-established harmony contained in the first organic germ, which, after innumerable transformations of the organic form and after millions of years, gave rise in the midst of the Amazonian region to an inedible Heliconide with certain yellow, black, and white markings on the wings, and at precisely the same time developed the tendency in a Pieride at the same spot on the globe to imitate this Heliconide as a model!

In addition to this assumption, which is certainly but little worthy of consideration, there is perhaps one other remaining, viz., that all or many Pierides and other species of butterflies possessed the same tendency to a Heliconoid variation and were always everywhere striving to develop this type, but succeeded only where they accidentally coincided in time and place with the model, the “tendency” being thus furthered by natural selection. But the facts negative this assumption, since such imitative variations have never been observed to a perceptible extent in other species.[295]

All variations which are demonstrably useful can be similarly dealt with if their origin is explained by variational tendencies.

We perceive that the objection which Von Hartmann brings against heredity is only valid on the ground that this process affords no security for the preservation of variations which occur singly. That heredity itself is a mechanical process is not directly disputed; it is simply assumed that new characters can be transferred by inheritance only when they are produced by the metaphysical “developmental principle,” and not when they arise “accidentally.” This critic does not therefore direct his attack against heredity, but rather against the mechanical origin of variability.

Von Hartmann might have said here that a reference of the phenomenon of heredity to purely mechanical causes, i.e. a mechanical theory of heredity, is up to the present time wanting. That he has not done so proves on the one hand that he despised the dialectical art, but, on the other hand, that he himself has not overlooked the subserviency of the total phenomenon to law, and that he grants the possibility of finding a mechanical explanation therefor. If, in fact, the power of inheritance does not depend upon mechanical principles, I know not what organic processes we are entitled to regard as mechanical, since they are all dependent in essence upon heredity, with which process they are at one, and from which they cannot be thought of as isolated. Haeckel correctly designates reproduction as surplus individual growth, and accordingly refers the phenomena of heredity to those of growth. Conversely, growth may also be designated reproduction, since it depends upon a continuous process of multiplication of the cells composing the organism, from the germ-cell to the innumerable congeries of variously differentiated cells of the highly developed animal body. Who can fail to see that these two processes, the reproduction of the germ-cell and its offspring in the economy of the individual, and the reproduction of individuals and species in the economy of the organic world, show an exact and by no means simply superficial analogy?[296] But whoso grants this must also conceive both processes to depend upon the same cause—he cannot assume for the one a causal power and for the other a directive principle. If nutrition and cell-multiplication are purely mechanical processes, so also is heredity. Although it has not yet been possible to demonstrate the mechanism of this phenomenon, it can nevertheless be seen broadly that by means of a minimum of living organic matter (e.g. the protoplasm of the sperm and germ-cell) certain motions are transferred, and these can be regarded as directions of development, as I have already briefly laid down in a former work.[297] The power of organisms to transmit their properties to their offspring appears to me to be only conceivable in such a manner “that the germ of the organism by its chemico-physical composition together with its molecular structure, has communicated to it a fixed direction of development—the same direction of development as that originally possessed by the parental organism....” (loc. cit. p. 24). This is confessedly nothing more than a hint, and we do not learn therefrom the means by which developmental direction can be possibly transferred to another organism.

Recently Haeckel, that indefatigable pioneer to whom we are indebted for such a rich store of new ideas, has attempted to bridge over this gap in his essay on “The Perigenesis of the Plastidule,” Berlin, 1876. The basic idea, that heredity depends upon the transference of motion, and variability upon a change of this motion, completely corresponds with the conviction gained in the province of physical science, that “all laws must finally be merged in laws of motion” (Helmholtz[298]). I hold this view to be the more completely justifiable—although certainly not in the remotest degree as proved—because I formerly designated the acquired individual variations as the “diversion of the inherited direction of development.” Haeckel’s hypothesis in so far accomplishes more than Darwin’s pangenesis, in which a transference of matter, and not of a species of motion peculiar to this matter, is assumed. But although the germ of a mechanical theory of heredity may be contained in Haeckel’s hypothesis, this nevertheless appears to me to be somewhat remote from completely solving the problem. It brings well into prominence one portion of the process of inheritance; under the image of a molecular motion of the plastidule, which motion is modifiable by external influences, we can well understand the fact of a change gradually taking place in the course of generations. On the other hand, the assumption of consciousness in the plastidule,—however admissible philosophically—although only as a formula, scarcely furnishes any deeper knowledge. In the light of a theory, detailed instances which were formerly obscure should become comprehensible. I fail to see, however, how the various forms of atavism, e.g. the reversions which so commonly occur by crossing different races, become more comprehensible by assuming consciousness in the plastidule. If in both parents the plastidule long ago acquired different molecular motions, why, in its rencounters in the germ, does it recollect past times and reassume the older and long abandoned motion? That it does acquire the latter is indeed a fact if we once refer the directional development of the individual to molecular motion of the plastidule; the wherefore does not appear to me, however, to become clearer by assuming consciousness in the plastidule. A mechanical theory of heredity must rather be able to show that the plastidule movements of the male and female germ-cells, in their rencounter in the case of the crossing of widely divergent forms, become mutually modified in such a manner that the motion of the common ancestral form must occur as the resultant. To such demonstration there is however as yet a long step. Haeckel himself moreover points out that his hypothesis is by no means a “mechanical theory of heredity,” but only an introduction to this theory, which he hopes “will be capable of being elevated to the rank of a genetic molecular theory” (loc. cit. p. 17). But although we must also confess with the critic of the “Philosophy of the Unconscious,” that “the facts of heredity have hitherto defied every scientific explanation,”[299] this furnishes us with no excuse for flying to a metaphysical explanation, “which is here certainly least able to satisfy the inability to understand the connection arising from natural laws.”

It is not to be wondered at that Von Hartmann, on the ground of the “Unconscious” on which he takes his stand, speaks of the law of correlation as an unconscious acknowledgment of a “non-mechanical universal principle on the side of Darwinism.” By “correlation” he understands something quite different to the idea which we attach to this expression. He supposes that “Darwinism sees itself compelled to acknowledge through empirical facts the uniform correlation of characters pertaining to the specific type; but it thereby contradicts its mechanical principles of explanation, all of which amount to the same thing as conceiving the type as a mosaic, chequered, superficial, and accidental aggregate of characters, which have been singly acquired, contemporaneously or successively, by selection or habit.” I do not believe, however, that any such conception has ever been admitted either by Darwin or any one else. The admission that not all, but only every deep-seated physiological detailed modification, is or may be bound up with a system of correlated changes, indeed implies that we on our side also acknowledge an internal harmony of parts—an equilibrium, as I have above expressed it.

But does this include the admission of a teleological principle, or exclude a mechanical explanation? Do we thereby acknowledge a “specific type” in the sense of an inseparably connected complex of characters, none of which can be taken away without all the others becoming modified? Does such a view agree generally with the empirical facts?

Neither of these views appears to me to represent the case.

I will first answer the second question. On all possible sides the earlier view of the absolute nature of species is contradicted; there is no boundary between species and varieties. But when Von Hartmann assumes that by the transformation of one species “into another” the “whole uniformly connected complex must become changed,” he falls back into the old doctrine of the absolute nature of species, which is sharply contradicted by multitudes of facts. We not unfrequently observe varieties which differ from the parent-form by only a single character, whilst others show numerous differences, and again others may be seen in which the differences predominate. This last deviation would then be designated by many systematists as a new species, but not so by others.

The “specific type” is thus indeed a kind of mosaic-work, but it is a structure to which all the single characters—the stones of the mosaic—belong and build up one harmonious whole, and not a meaningless confusion. Some of the stones or groups of stones can be taken away and replaced by others differently coloured without the structure being thereby necessarily distorted, i.e. destroyed as a structure; but the larger the stones which are exchanged the more necessary will corrections in the other parts of the structure become, in order that the harmony of the whole may be preserved.

Still more weighty than those insensible transitions which in various groups of animals so frequently connect species with species, appear to me, however, the facts made known in the second essay of the second part of this volume, which prove that the two forms in which one species appears can change entirely independently of one another. The caterpillar changes and becomes a new variety or even species (according to the form-value of the change), whilst the butterfly remains unaltered. How could this occur if some other law than that of physiological equilibrium linked together the parts or characters and permitted them to become severed? Must not the two stages become changed with and through one another, like the parts of one body, since they first together constitute the specific type? Is not the fact of this not happening a proof that the whole “uniformly connected complex” of the specific type is not bound and held together by a metaphysical principle, but simply by natural laws?

Now when Von Hartmann comprises the relations of different species to one another under the idea of correlation, such for instance as the relation of dependence in which orchidaceous flowers stand with respect to the insects which visit them, he completely abandons the scientific conception which should be associated with this expression, and compares together two heterogeneous things which have nothing in common excepting that they are both considered by him as a result of the “Unconscious.” The consequence which is then deduced from this correlation of his own construction, viz., that an organic law of correlation is only another expression for a “law of organic development” in the sense of a metaphysical power, obviously cannot be admitted.

By correlation we understand nothing more than the dependence of one part of the organism upon the others and the mutual inter-relations of these parts, which depend entirely upon a “physiological relation of dependence,” as Von Hartmann himself has correctly designated it. Herein is evidently comprised the total morphology of the organism—the structure as a whole, the length, thickness and weight of the single parts, as well as the histological structure of the tissues, since upon all these depends the performance of the single parts. But when, under correlation, Von Hartmann comprises “also a morphological, systematic, inter-action of all the elements of the organism with reference both to the typical ground-plan of the organization as well as to the microscopic anatomical structure of the tissues,” he drags into the idea something foreign to it, not on the ground of facts, but actually in opposition to them, and supported only by a supposed “innate developmental principle” which “is not of a mechanical nature.”

The living organism has already been often compared with a crystal, and the comparison is, mutatis mutandis, justifiable. As in the growing crystal the single molecules cannot become joined together at pleasure, but only in a fixed manner, so are the parts of an organism governed in their respective distribution. In the crystal where nothing but homogeneous parts become grouped together their resulting combination is likewise homogeneous, and it is obvious that they offer but very little possibility of modification, so that the governing laws thus appear restricted and immutable. In the organism, whether regarded microscopically or macroscopically, various parts become combined, and these therefore offer numerous possibilities of modification, so that the governing laws are more complex, and appear less restricted and unchangeable. In neither instance do we know the final causes which always lead to a given state of equilibrium; in the case of a crystal it has not occurred to anybody to ascribe the harmonious disposition of the parts to a teleological power; why then should we assume such a force in the organism, and thus discontinue the attempt, which has already been commenced, to refer to its natural causes that harmony of parts which is here certainly present and equally conformable to law?

On these grounds the assertion that the theory of selection is not an attempt at a “mechanical” explanation of organic development appears to me to be incorrect. Variability and heredity, as well as correlation, admit of being conceived as purely mechanical, and must be thus regarded so long as no more cogent reasons can be adduced for believing that some force other than physico-chemical lies concealed therein.

But we certainly cannot remain at the purely empirical conception as laid down by Darwin in his admirable work on the “Origin of Species.” If the theory of selection is to furnish a method of mechanical explanation, it is essential that its factors should be formulated in a precise mechanical sense. But as soon as we attempt to do this it is seen that, in the first enthusiasm over the newly discovered principle of selection, the one factor of transformation contained in this principle itself has been unduly pushed into the background, to make way for the other more apparent and better known factors.

I have for many years insisted that the first, and perhaps most important, or in any case the most indispensable, factor in every transformation, is the physical nature of the organism itself.[300]

It would be an error to believe that it is entirely the external conditions which determine what changes shall appear in a given species; the nature of these changes depends essentially upon the physical constitution of the species itself, and a modification actually arising can obviously be only regarded as the resultant of this constitution and of the external influences acting thereon.

But if an essential or perhaps even a preponderating share in determining new characters is to be undoubtedly ascribed to the organism itself, for a mechanical representation of organic developmental processes everything depends upon our being able to conceive this most important factor in a definite theoretical manner, and to comprise under one common point of view its apparently contradictory manifestations of constancy and variability.

Now every change of considerable extent is certainly considered by Darwin to be the direct or indirect consequence of external actions; but indirect action always presupposes a certain small variability (individual variability), without which larger modifications cannot be brought about. Empirically this small amount of variability is doubtless present, but the question arises, upon what does it depend? Can it be conceived as arising mechanically, or is it perhaps just at this point that the metaphysical principle steps in and offers those minute variations which make possible that course of development which, according to this view, is immutably pre-determined? It is certainly the absence of a theoretical definition of variability which always leaves open a door for smuggling in a teleological power. A mechanical explanation of variability must form the basis of this side of the theory of selection.

This explanation is not difficult to find. All dissimilarities of organisms must depend upon the individuals having been affected by dissimilar external influences during the course of the development of organic nature. If we ascribe to the organism the power of giving rise by multiplication only to exact copies of itself, or, more correctly, the power of transmitting unaltered to its successors the motion of its own course of development, each “individual variation” must depend upon the power of the organism to react upon external influences, i.e. to respond by changes of form and of function, and consequently to modify its original (inherited) developmental direction.

It has sometimes been insisted upon, that the “individuals of the same species” or the offspring of one mother cannot be absolutely equal, because, from the commencement of their existence, they have been subjected to dissimilar actions of the environment. But this implies that by perfectly equal influences they would become equal, i.e. it supposes that variability is not inseparably bound up with the essence of the organism, but is only the consequence of developmental tendencies which are in themselves equal being unequally influenced. As a matter of fact the first germs of an individual certainly cannot be supposed to be perfectly equal, because the individual differences of the ancestors must be contained therein in different degrees according to their constitution, and we should have to go back to the primordial organism of the earth in order to find a perfectly homogeneous root, a tabula rasa from which the descendants would commence their development. Whether such a homogeneous root ever existed is however doubtful; it is much more probable that numerous organisms first arose spontaneously,[301] and these cannot be presumed to have been absolutely equal, since the conditions under which they came into life cannot have been perfectly identical. Let us, however, for the sake of simplicity assume a single primordial organism; the first generation which took its rise from this by reproduction could only have possessed such individual differences as were produced by the action of dissimilar external influences. But the third generation, together with self-acquired, would also have shown inherited, dissimilarities, and in each succeeding generation the number of tendencies to individual difference imparted to the germ by heredity must have increased to a certain degree, so that it may be said that all germs, from their first origination, bear in themselves a tendency to show individual peculiarities, and would develop these even if they should not be again affected by dissimilar influences. This is obviously the case, since the youngest egg-cells in the ovary of an animal are, as can be demonstrated, always exposed to unequal external conditions with respect to nutrition and pressure.[302] Hence, if it were possible that two germs were exactly equal with respect to the direction of development imparted to them by heredity, they would nevertheless furnish two incongruent individuals; and if, conversely, it were possible that two individuals could be exposed to absolutely the same external influences from the formation of the embryo, these also could not be identical, because the individual differences of the ancestors would entail small differences, even in asexual reproduction, in the direction of development transmitted to the egg. The differences between individuals of similar origin thus finally depend entirely upon the dissimilarity of external influences—on the one side upon those which divert the development of the progenitors, and on the other side upon those which divert the individual itself from its course, i.e. from the developmental direction transmitted hereditarily. Although I thus essentially agree with Darwin and Haeckel in so far as these authors refer the “universal individual dissimilarity” to dissimilar external actions, I differ from Darwin in this, that I do not see an essential distinction between the direct and indirect production of individual differences, if by the latter is meant only the unequal influencing of the germ in the parental organism. Haeckel is certainly correct in referring the “primitive differences of the germs produced by the parents” to the inequalities of nutrition to which the single germs must inevitably have been exposed in the parent organism; but another dissimilarity of the germs must evidently be added—a dissimilarity which has nothing to do with unequal nutrition, but which depends upon unequal inheritance of the individual differences of the ancestors, a source of dissimilarity which must arise to a greater extent in sexual than in asexual reproduction. Just as in sexual propagation there occurs a blending of the characters (or more precisely, developmental directions) of two contemporaneous individuals in one germ, so in every mode of reproduction there meet together in the same germ the characters of a whole succession of individuals (the ancestral series), of which the most remote certainly make themselves but seldom felt in a marked degree.

The fact of individual variability can in this way be well understood; the living organism contains in itself no principle of variability—it is the statical element in the developmental processes of the organic world, and would always reproduce exact copies of itself if the inequality of the external influences did not affect the developmental course of each new individual; these influences are therefore the dynamical elements of the process.

From this conception of variability two important empirically established facts can be theoretically deduced, viz. the limitability of variation with respect to quality, which has already been previously mentioned, and the origination of transformations by the direct action of external conditions of life.

If the differences in individuals of the same origin depend upon the action of unequal influences, variation itself is nothing else than the reaction of the organism to a definite external inciting cause, the quality of the variation being determined by the quality of the inciting cause and by that of the organism. In the cases of individual variation hitherto considered, the quality of the organism is equal but that of the inciting cause is unequal, and in this way there arise minute differences in organisms of an equal physical constitution—variations of a different quality.

The same result, viz., different qualities of variation, may also arise in a reverse manner by organisms of a different physical nature being affected by equal external influences. The response of the organism to the cause inciting change would be different according to its nature, or, in other words, organisms of different natures react differently when affected by equal modifying influences. The physical nature of the organism plays the chief part with respect to the quality of the variations; each specific organism can thus give rise to extremely numerous, but not to all conceivable, variations; that is, only to such variations as are made possible by its physical composition. From this it follows further that the possibilities of variation in two species are more widely different, the wider they diverge in physical constitution (including bodily morphology)—that a cycle of variation is peculiar to every species. In this manner we are led to the knowledge that there must certainly exist a “fixed direction of variation,” but not in the sense of Askenasy and Von Hartmann, as the result of an unknown internal principle of development, but as the necessary, i.e. mechanical, consequence of the unequal physical nature of the species, which must respond even to the same inciting cause by unequal variations.

The facts, as far as we know them, agree very well with this conclusion. Allied species vary in a similar manner, whilst species which are more distantly related vary in a different manner, even when acted upon by the same external influences. Thus, in the first part of these “Studies” I have remarked that many butterflies under the influence of a warm climate acquire an almost black coloration (Polyommatus Phlæas), whilst on the other hand others become lighter (Papilio Podalirius).

We can thus understand why always certain courses of development are followed, a fact which cannot be completely explained by the nature of the conditions of life which induce the variations. But as soon as we clearly perceive that the quality of the changes essentially depends upon the physical nature of the organism itself, we arrive at the conclusion that species of widely diverging constitutions must give rise to different variations, whilst those of allied constitutions would produce similar variations. But definite courses of development are thus traced out, and we perceive that from any point of the organic developmental series, it is impossible that any other point can be attained at pleasure. Variation in a definite direction thus by no means necessitates the acknowledgment of a metaphysical developmental principle, but can be well conceived as the mechanical result of the physical constitution of the organism.

The manner in which the dissimilar physical constitution of organisms must arise can also be easily shown, although the first commencement of the whole developmental series, i.e. the oldest living forms must be assumed to have been almost homogeneous in their physical constitution. The quality of the variation is, as said before, not merely the product of the physical constitution, but the resultant of this and of the quality of the changing external conditions. Thus from the first “species” there proceeded, through the dissimilar influence of external conditions of life, several new “species,” and as this took place the former physical nature of the organism at the same time became changed, necessitating also a new mode of reacting upon external influences, i.e. another direction of variation. The difference from the primary “species” must certainly be conceived as having been very minute, but it must have increased with each new transformation, and must have proceeded exactly parallel with the degree of physical change connected with each transformation. Thus, hand in hand with the modifications, the power of modification, or mode of reaction of the organism to changing influences, must have continually become re-modified, and we finally obtain an endless number of differently constituted living forms, of which the variational tendencies are different in exact proportion to their physical divergence, so that nearly allied forms respond similarly, and widely divergent forms very differently, to the same inciting causes.

Individual variation arises, as I have attempted to show, by each individual having been continually affected by different, and indeed by constantly changing, influences. Let us, however, imagine on the contrary, that a large group of individuals is affected by the same influences—in fact by such influences as the remaining individuals of the species are not exposed to: this group of individuals would then vary in a nearly similar manner, since both factors of variation, viz. the external influence and the physical constitution, are equal or nearly so. Such local variations would first become prominent when the same external influence had acted upon a series of generations, and the minima of variation produced in the individual by the once-exerted action of the cause inciting change had become augmented by heredity. Transformations of some importance (up to the form-value of species) can thus arise simply by the direct action of the environment, in the same way as that in which individual differences are produced—only the latter fluctuate from generation to generation, since the inciting influences continually change; whilst, in the former, the constant external cause inciting modification always reproduces the same variation, so that an accumulation of the latter can take place. Climatic varieties can be thus explained.

A more efficacious augmentation of the variations arising in the single individual is certainly brought about by the indirect action of the environment upon the organism. It is not here my intention to explain once more the processes of natural selection; I mention this only in order to point out that in these cases transformation depends upon a double action of the environment, since the latter first induces small deviations in the organism by direct action, and then accumulates by selection the variations thus produced.

By regarding variability in this manner—by considering each variation as the reaction of the organism to an external action, as a diversion of the inherited developmental direction, it follows that without a change in the environment no advance in the development of organic forms can take place. If we imagine that from any period in the earth’s history the conditions of life remain completely unchanged, the species present on the earth at this period would not, according to our view, undergo any further modification. Herein is clearly expressed the difference of this view from that other one according to which the inciting principle of modification is not in the environment, but lies in the organism itself in the form of a phyletic vital force.

I cannot here refrain from once more returning to the old (ontogenetic) vital force of the natural philosophers, since the parallel between this and its younger sister, the “phyletic vital force” which appears in so many disguises, is indeed striking. Were the inciting principle of the development of the individual actually an independent vital force acting within the organism, the birth and growth of the individual would be able to take place without the continuous encroachment of the environment, such as occurs in nutrition and respiration. Now this is known to be impossible, so that those who support the existence of such a force, if any still exist, would be driven to the obscure idea of a co-operation between the designing power and the influences of the environment, just in the same manner as such a co-operation is at present postulated by the defenders of the phyletic vital force. I shall further on take the opportunity of pointing out that this last idea is quite untenable; with respect to the (ontogenetic) vital force any clearer proof cannot well be adduced, but it will be admitted that the confused notion of the co-operation and inter-action of teleological and causal powers is, from our point of view, opposed to those very simple and clear ideas which are in harmony with the views on phyletic development. As in racial development each change of the organic type is entirely dependent upon the action of the environment upon the organism, so in the development of the individual, the totality of the phenomena of the personal life must depend upon similar actions. Physiology, as is known, herein entirely supports our view, since this shows that without the continual alternating action of the environment and of the organism there can be no life, and that vital phenomena are nothing but the reactions of the organism to the influences of the environment.

It will be immediately perceived how exactly the processes of phyletic and of ontogenetic development coincide, not merely in their external phenomena but in their nature, if we trace the consequences of the existing knowledge of the structure of the animal body. Although we may not entirely agree with Haeckel’s doctrine of individuality in its details, its correctness must on the whole be conceded, since it cannot be disputed that the notion of individuality is a relative one, and that several categories of morphological individuals exist, which appear not only singly as physiological individuals, i.e. as independent living beings of lowest grade, but which can also combine to form beings of a higher order.

But if we admit this, we should see with Haeckel nothing but reproduction in the origination of a high organism from a single cell, the egg; this reproduction being at the same time combined with various differentiations of the offspring, i.e. with adaptations of the latter to various conditions of life. Not even in the fact that the tissues and organs of a single physiological individual stand in great dependence upon one another through physical causes,[303] is there any striking difference between this view and the phyletic composition of the animal (and vegetable) kingdom out of physiological individuals (Haeckel’s “Bionten”), since contemporaneous animals (individuals and species) are known to influence one another in the most active manner.

Now if we further consider that the same units (cells) which, by their reproduction and division of labour, at present compose the body of the highest organism, must at one time have constituted as independent beings the beginning of the whole of organic creation, and that consequently the same processes (division of cells) which now lead to the formation of a mammal, at that time led only to a long series of different independent beings, it will be admitted that both developmental series must depend upon the same inciting powers, and that with reference to the causes of the phenomena it is not possible that any great gap can exist between ontogeny and phylogeny, i.e. between the life-phenomena of the individual and those of the type. According to our view both depend upon that co-operation of the same material physical forces which admits of being briefly summarized as the reaction of organized living matter to influences of the environment.

Our opponents either cannot boast of such harmony in their conception of nature, or else they must, together with the phyletic vital force, re-admit into their theory the old ontogenetic vital force. I know not indeed why they should not do so. Whoever inclines to the view that organic nature is governed not merely by causal, but at the same time by teleological, forces, may admit that the latter are as effective as inciting causes of individual, as they are of phyletic, development. According to my idea they are even bound to admit this, since it cannot be perceived why the adaptations of the ontogeny should not depend upon the same metaphysical principle assumed for each individual, as the adaptations of the phylogeny; the latter are indeed only brought about by the former. I believe therefore that the vital force (ontogenetic) of the ancients stands or falls with the modern (phyletic) vital force. We must admit both or neither, since they both rest on the same basis, and are supported or opposed by the same arguments. Whoever feels justified in setting up a metaphysical principle where complete proof that known forces are sufficient for the explanation of the phenomena has not yet been adduced, must do the same with respect to individual, as he does to phyletic, development, since this proof is in both cases very far from being complete, and still contains large and numerous gaps.[304]

The theoretical conception of variation as the reaction of the organism to external influences has also not yet been experimentally shown to be correct. Our experiments are still too coarse as compared with the fine distinctions which separate one individual from another; and the difficulty of obtaining clear results is greatly increased by the circumstance that a portion of the individual deviations always depends upon heredity, so that it is frequently not only difficult, but absolutely impossible, to separate those which are inherited from those which are acquired. Still further are we removed from being able to refer variation to its final mechanical causes, i.e. from a mechanical theory of reproduction, which would bring within the range of mathematical calculation both the phenomena of stability (heredity) and of change (variability).

But although sufficient proofs of the correctness of the views here advocated cannot at present be adduced, these views are not contradicted by any known facts—they are, on the contrary, supported by many facts which they in turn make comprehensible (local forms, different cycles of variation in heterogeneous species). These views are finally completely justified by their furnishing the only possible theoretical formulation of variability on which a mechanical conception of organic development can be based. That such a conception is not only admissible, but is unavoidable, at least to the naturalist, I have already attempted to prove.

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II.
Mechanism and Teleology.

In the third volume of his smaller works Karl Ernst von Baer submits the theory of selection to a most searching examination. Without actually calling in question its scientific admissibility, he believes that this theory is dependent upon its satisfying one condition, viz. that it should connect the teleological with the mechanical principle.

“The Darwinian hypothesis, as stated by its supporters, always ends in denying to the processes of nature any relation to a future, i.e. any relation of aim or design. Since such relations appear to me quite evident,” &c. And further:—“If the scientific correctness of the Darwinian hypothesis is to be admitted, it must accommodate itself to this universal striving after a purpose. If it cannot do this we should have to deny its value.”

These words appear almost equivalent to passing a sentence of doom upon the theory of selection and the mechanical conception of nature, for how can one and the same process be effected simultaneously by necessity and by designing powers? The one excludes the other, and we must—so it appears—take our stand either on one side or the other.

Nevertheless we cannot set aside Von Baer’s proposition without further examination simply because it is apparently incapable of being fulfilled, since it contains a truth which should not be overlooked, even by those who uphold the mechanical theory of nature. It is the same truth which is also made use of by the philosophical opponents of this theory, viz. that the universe as a whole cannot be conceived as having arisen from blind necessity—that the endless harmony revealed in every nook and corner by all the phenomena of organic and of inorganic nature cannot possibly be regarded as the work of chance, but rather as the result of a “vast designed process of development.” It is also quite correct when, in reply to the supposed objection that the mechanical theory of nature is not concerned with chances but with necessities, Von Baer answers that the operations of a series of necessities which “are not connected together” can only be termed accidents in their opposing relations. He illustrates this by instancing a target. If I hit the latter by a well-aimed shot, nobody would explain this as the result of an accident, but if “a horseman is riding along a gravelly road past this target, and one of the pebbles thrown up by the hoof of the galloping horse hits the mark, this would be termed an accident of extremely rare occurrence. My target was not the mark for the pebble, therefore the hit was purely accidental, although the projection of the stone in this precise direction with the velocity which it had acquired, was sufficiently explained by the kick given by the horse. But the hit was accidental because the kick of the galloping horse, although it necessarily projected the pebble, had no relation at all to my target. For the same reason we must regard the universe as an immense accident if the forces which move it are not designedly regulated—the more immense because it is not a single motion of projection that acts here, but a large number of heterogeneous powers, i.e. a large number of variously acting necessities which are, as a whole, devoid of purpose, but which nevertheless accomplish this purpose, not only at any single moment, but constantly. A truly admirable series of desirable accidents!”[305]

The same idea is expressed, although in a very different manner, by Von Hartmann, in the concluding chapter of his work already quoted. He thinks that “design is a necessary and certain consequence of the mechanical laws of nature.” “Were the mechanism of natural laws not teleological there would be no mechanically regulated laws, but a weak chaos of obstinate and capricious powers. Not until the causality of the laws of inorganic nature had superseded the expression “dead” nature, and had shown itself as the mainspring of life and of a conformability to design visible on all sides, did it deserve the name of mechanical lawfulness; just as a complication of wheels and machinery made by man, which move in some definite manner with respect to one another, only acquires the name of a mechanism or of a machine when the immanent teleology of the combination and of the various movements of the parts is revealed.”[306]

Against the correctness of the idea underlying these statements scarcely anything can in my opinion be said. The harmony of the universe and of that portion of it which we designate organic nature, cannot be explained by chance, i.e. without a common ground for co-operating necessities; by the side of mere mechanism it is impossible not to acknowledge a teleological principle—the only question is, in what manner can we conceive this as acting without abandoning the purely mechanical conception of nature?

This is obviously effected if, with Von Baer and Von Hartmann, we permit the metaphysical principle to interrupt the course of the mechanism of nature, and if we consider both the former and the latter to work together with equal power. Von Hartmann expressly makes such an admission under the name of an “internal principle of development,” to which he attributes such an important share that one cannot understand why it should have any need for the employment of causal powers, and why it does not simply do everything itself. Von Baer expresses himself much less decisively, and even in many places insists upon the purely mechanical connection of organic natural phenomena; but that with him also the idea of interruption by a metaphysical principle is present, is principally shown by his assuming, at least partly, the per saltum development of species. This necessarily involves an actively internal power of development.

Although I have already brought forward many arguments against the existence of such a power, and although in refuting it every form of development by directive powers is at the same time overthrown, it nevertheless appears to me not to be superfluous in such a deeply important question to show that a per saltum development, and especially the so-called heterogeneous generation, is inconceivable, not only on the ground of the arguments formerly employed against the phyletic vital force in general, but quite independently of these.

In the first place it must be said that the positive basis of this hypothesis is insecure. Cases of sudden transformation of the whole organism with subsequent inheritance are as yet quite unknown. It has been shown that the occasional transformation of the Axolotl must most probably be regarded in a different light. Another case, taken for heterogeneous generation, viz. the budding of twelve-rayed Medusæ in the gastric cavity of an eight-rayed species, has lately been shown by Franz Eilhard Schulze[307] to be a kind of parasitism or commensalism. The buds of the Cuninæ do not spring, as was supposed, from the Geryonia, but are developed from a Cunina egg. But even if we recall here the cases of alternation of generation and heterogenesis, this would not be of any value by way of proof; it would only be thus indicated how one might picture to oneself a sudden transformation. That in alternation of generation, or generally, in every mode of cyclical reproduction, we have not to deal with the abandonment of one type of organization and the transition to some other, is proved by the continual return to the type of departure—by the cyclical character of the entire transformation. That two quite heterogeneous types can belong to one cycle of development is, however, capable of a far better and more correct explanation than would be given by the supporters of per saltum development. If we trace cyclical reproduction to the adaptation of different developmental stages or generations to deviating conditions of life, we thus not only explain the exact and often striking agreement between form and mode of life—we not only bridge over the gap between metamorphosis and alternation of generation, but we can also understand how, within one and the same family of Hydrozoa, species can occur with or without alternation of generation, and further how other species can exist in which the alternation of generation (the production of free Medusæ) is limited to the one sex; we can understand in general how one continuous series of forms may lead from the simple sexual organ of the Polypes to the independent and free swimming sexual form of the Medusæ, and how hand in hand with this the simple reproduction becomes gradually cyclical. It is just these intermediate steps between the two kinds of reproduction that make quite untenable the idea that the heterogeneous forms in cyclical propagation arise through so-called “heterogeneous generation,” i.e. through sudden per saltum transformation. It is excusable if philosophers to whom these facts are strange, or who have to take the trouble of working them up, should adduce alternation of generation as an instance of “heterogeneous generation,” but by naturalists this should be once and for ever abandoned.

All other facts which have hitherto been referred to “heterogeneous generation” are still less explicable as such, inasmuch as they always relate to changes in single parts of an organism, such as the sudden change of fruit or flower in cultivated plants. The notion of per saltum development, however, demands a total transformation—it comprises (as Von Hartmann quite correctly and logically admits) the idea of a fixed specific type which can only be re-modelled as a whole, and cannot become modified piecemeal. It must further be added, that the observed variations which have arisen abruptly in single parts are not as a rule inherited:[308] fruit-trees are only propagated by grafting, i.e. by perpetuating the individual, and not by ordinary reproduction by seeds. Now, if we nowhere see sudden variations of large amount perpetuated by heredity, whilst we everywhere observe small variations which can all be inherited, must it not be concluded that per saltum modification is not the means which Nature employs in transforming species, but that an accumulation of small variations takes place, these leading in time to large differences? Is it logical to reject the latter conclusion because our period of observation is too brief to enable us to directly follow long series of accumulations, whilst per saltum variation is admitted, although unsupported by a single observation? As long as there remains any prospect of tracing large deviations to the continually observed phenomenon of small variations, I believe we have no right to resort to the purely hypothetical explanation afforded by per saltum variations.

But the hypothesis of “heterogeneous generation” is not only without a basis of facts—it can also be directly shown to be untenable. Since the operation of an internal power of transformation does not explain adaptation to the conditions of life, the claims of natural selection to explain these transformations must be admitted; but the co-operation of a phyletic vital force and natural selection is inconceivable if we imagine the modifications to occur per saltum.

The supposed “heterogeneous generation” is always illustrated by the example of alternation of generation. The origination of a new animal form is thus conceived to take place in the same manner as we now see, in the cyclical reproduction of the Medusæ, free swimming, bell-shaped Medusoids, produced from fixed polypites, or Cercariæ from Trematode worms by internal budding; in brief, it is imagined that one animal form suddenly gives rise to another widely deviating form by purely internal causes. Now on this theory it would be an unavoidable postulate, that by such a process of per saltum development there arises not merely a new type of some species, but at the same time individuals capable of living and of persisting under, and fitted to, given conditions of life. But every naturalist who has attempted to completely explain the relation between structure and mode of life knows that even the small differences which separate one species from another, always comprise a number of minute structural deviations which are related to well defined conditions of life—he knows that in every species of animal the whole structure is adapted in the most exact manner in every detail to special conditions of life. It is not an exaggeration when I say in every detail, since the so-called “purely morphological parts” could not be other than they are without causing changes in other parts which exercise a definite function. I will not indeed assert that in the most closely related species all the parts of the body must in some manner differ from one another, if only to a small extent; it seems to me not improbable, however, that an exact comparison would very frequently give this result. That animals which are so widely removed in their morphological relations as Medusæ and Polypes, or Trematoda and their “nurses,” are differently constructed in each of their parts can, however, be stated with certainty.

Now if this wide deviation in every part were in itself no obstacle to the assumption of a designing and re-modelling power, it would become so by the circumstance that all the parts of the organism must stand in the most precise relation to the external conditions of life, if the organism is to be capable of existing—all the parts must be exactly adapted to certain conditions of life. How can this be brought about by a transforming force acting spasmodically? Von Hartmann—who, in spite of his clear perception and widely extended scientific knowledge, cannot possibly possess a strong conviction of that harmony between structure and life-conditions prevailing throughout the whole system of the organism, and which personal research and contemplation are alone able to give—simply bridges over the difficulty by permitting natural selection to come to his aid as an “auxiliary principle” of the re-modelling power. It would not be supposed that naturalists would resort to the same device—nevertheless those who support the phyletic force and per saltum development generally invoke natural selection as the principle which governs adaptation. But when does this agency come into operation? When by germinal metamorphosis a new form has arisen, this, from the first moment of its existence, must be adapted to the new conditions of life or it must perish. No time is allowed for it to continue in an unadapted state throughout a series of generations until adaptation is luckily reached through natural selection. Let us have either natural selection or a phyletic force—both together are inconceivable. If there exists a phyletic force, then it must itself bring about adaptation.

It might perhaps be here suggested that the same objection applies to that process of modification which is effected by small steps, but that it does so only when the change occurs suddenly. This, however, as I have already attempted to show, but very rarely takes place; in many cases (mimicry) the conditions even change in the first place through the change in form and therefore, as is evident, as gradually as the latter. It must be the same in all other cases where transformation of the existing form and not merely extinction of the species concerned takes place. The transmutation must always keep pace with the change in the conditions of life, since if the latter change more rapidly the species could not compete with rival species—it would become extinct.

The abrupt transformation of species implies sudden change in the conditions of life, since a Medusa does not live like a Polype, nor a Trematode like its “nurse.” For this reason it is impossible that natural selection can be an aiding principle of “heterogeneous generation.” If such abrupt transformation takes place it must produce the new form instantly equipped for the struggle for existence, and adapted in all its organs and systems of organs to the special conditions of its new life. But would not this be “pure magic”? It is not thereby even taken into consideration that here—as in the cases of mimicry—time and place must agree. The requirements of a pre-established harmony (“prästabilirte Harmonie”) further demand that an animal fitted for special conditions of life should only make its appearance at that precise period of the earth’s history when these special conditions are all fulfilled, and so forth.

But he who has learnt to perceive the numerous and fine relations which, in every species of animal, bring the details of structure into harmony with function, and who keeps in view the impelling power of these conditions, cannot possibly hold to the idea of a per saltum development of animal forms. If development has taken place, it must have occurred gradually and by minute steps—in such a manner indeed that each modification had time to become equilibrated to the other parts, and in this way a succession of modifications gradually brought about the total transformation of the organism, and at the same time secured complete adaptation to new conditions of life.

Not only abrupt modification however, but every transformation is to be rejected when based upon the interference of a metaphysical principle of development. Those to whom the arguments already advanced against such a principle appear insufficient may once more be asked, how and where should this principle properly interfere? I am of opinion that one effect can have but one sufficient cause; if this suffices to produce it, no second cause is required. The hand of a watch necessarily turns once round in a circle in a given time as soon as the spring which sets the mechanism in movement is wound up; in an unwound watch a skilful finger can perhaps give the same movement to the hand, but it is impossible that the latter can receive both from the operator and from the spring at the same time, the same motion as that which it would receive through either of these two powers alone. In the same manner it appears to me that the variations which lead to transformation cannot be at the same time determined by physical and by metaphysical causes, but must depend upon either one or the other.

On no side will it be disputed that at least one portion of the processes of organic life depends upon the mechanical co-operation of physical forces. How is it conceivable that sudden pauses should occur in the course of these causal forces, and that a directive power should be substituted therefor, the latter subsequently making way again for the physical forces? To me this is as inconceivable as the idea that lightning is the electric discharge of a thunder-cloud, of which the formation and electrical tension depends upon causal forces, and of which the time and place are purely determined by such forces, but that Jupiter has it nevertheless in his power to direct the lightning flash according to his will on to the head of the guilty.

Now although I deny the possibility or conceivability of the contemporaneous co-operation of teleological and of causal forces in producing any effect, and although I maintain that a purely mechanical conception of the processes of nature is alone justifiable, I nevertheless believe that there is no occasion for this reason to renounce the existence of, or to disown, a directive power; only we must not imagine this to interfere directly in the mechanism of the universe, but to be rather behind the latter as the final cause of this mechanism.

Von Baer himself points this out to us, although he does not follow up the complete consequences of his arguments. He especially insists in his book, which abounds in beautiful and grand ideas, that the notions of necessity (causality) and of purpose by no means necessarily exclude one another, but rather that they can be connected together in a certain manner. Thus, the watchmaker attains his end, the watch, by combining the elastic force of a spring with wheel-work, i.e. by utilizing physical necessities; the farmer accomplishes his purpose, that of obtaining a crop of corn, by sowing the seed in suitable land, but the seed must germinate as an absolute necessity when exposed to the influences of warmth, soil, moisture, &c. Thus, in these instances a chain of necessities is undoubtedly connected with a teleological force, the human will; and it directly follows from such cases that wherever we see an aim or result attained through necessities, the directive force does not interrupt the course of the series of necessities which have already commenced, but is active before the first commencement of these necessities, since it combines and sets the latter in movement. From the moment when the mechanism of the watch is combined harmoniously and the spring wound up, it goes without the further interference of the watchmaker, just as the corn-seed when once placed in the earth develops into a plant without assistance from the farmer.

If we apply this argument to the development of the organic world, those who defend mechanical development will not be compelled to deny a teleological power, only they would have with Kant[309] to think of the latter in the only way in which it can be conceived, viz. as a Final Cause.

In the region of inorganic nature nobody any longer doubts the purely mechanical connection of the phenomena. Sunshine and rain do not now appear to us to be whims of a deity, but divine natural laws. As the knowledge of the processes of nature advances, the point where the divine power designedly interrupts these processes must be removed further back; or, as the author of the criticism of the philosophy of the Unconscious[310] expresses it, all advance in the knowledge of natural processes depends “upon the continual elimination of the idea of the miraculous.” We now believe that organic nature must be conceived as mechanical. But does it thereby follow that we must totally deny a final Universal Cause? Certainly not; it would be a great delusion if any one were to believe that he had arrived at a comprehension of the universe by tracing the phenomena of nature to mechanical principles. He would thereby forget that the assumption of eternal matter with its eternal laws by no means satisfies our intellectual need for causality. We require before everything an explanation of the fact that relationships everywhere exist between the parts of the universe—that atoms everywhere act upon one another.[311] He who can content himself with the assumption of matter may do so, but he will not be able to show that the assumption of a Universal Cause underlying the laws of nature is erroneous.

It will not be said that there is no advantage in assuming such a Final Cause, because we cannot conceive it, and indeed cannot so much as demonstrate it with certainty. It certainly lies beyond our power of conception, in the obscure region of metaphysics, and all attempts to approach it have never led to anything but an image or a formula. Nevertheless there is an advance in knowledge in the assumption of this Cause which well admits of comparison with those advances which have been led to by certain results of the new physiology of the senses. We now know that the images which give us our sense of the external world are not “actual representations having any degree of resemblance,”[312] but are only signs for certain qualities of the outer world, which do not exist as such in the latter, but belong entirely to our consciousness. Thus we know for certain that the world is not as we perceive it—that we cannot perceive “things in their essence”—and that the reality will always remain transcendental to us. But who will deny that in this knowledge there is a considerable advance, in spite of its being for the most part of a negative character? But just as we must assume behind the phenomenal world of our senses an actual world of the true nature of which we receive only an incomplete knowledge (i.e. a knowledge corresponding only in reality with the relations of time and space), so behind the co-operating forces of nature which “aim at a purpose” must we admit a Cause, which is no less inconceivable in its nature, and of which we can only say one thing with certainty, viz., that it must be teleological. Just as the former first leads us to perceive the true value of our sensual impressions, so does the latter knowledge lead us to foresee the true significance of the mechanism of the universe.

It is true that in neither case do we learn more than that there is something present which we do not perceive, but in both instances this knowledge is of the greatest value. The consciousness that behind that mechanism of the universe which is alone comprehensible to us there still lies an incomprehensible teleological Universal Cause, necessitates quite a different conception of the universe—a conception absolutely opposed to that of the materialist. Most correctly and beautifully does Von Baer say that “a purpose cannot be otherwise conceived by us than as proceeding from a will and consciousness; in this would the ‘aiming at a purpose,’ which appears to us as reasonable as it is necessary, have its deepest root.” If we conceive in this world a divine Universal Power exercising volition as the ultimate basis of matter and of the natural laws resident therein, we thus reconcile the apparent contradiction between the mechanical conception and teleology. In the same way that Von Hartmann, somewhere speaks of the immanent teleology of a machine, we might speak of the immanent teleology of the universe, because the single forces of matter are so exactly adjusted that they must give rise to the projected world, just as the wheels and levers of a machine bring forth a required manufactured article. I admit that these are grossly anthropomorphic ideas. But as mortals can we have any other ideas? Is not the notion of purpose in itself an equally anthropomorphic one? and is there any certainty that the idea of causality is less so? Do we know that causality is unlimited, or that it is universally valid? In the absence of this knowledge, should it not be permissible to satisfy as far as we can the craving of the human mind for a spiritual First Cause of the universe, by speaking of it in terms conceivable to human understanding? We can take up such a final position and still be conscious that we thereby form no certain conception, and indeed come no nearer to the reality. The materialist still makes use of the notion of “eternity,” and frequently handles it as though it were a perfectly known quantity. We nevertheless do not seriously believe that by the expression “eternal matter,” any true idea resulting from human experience is gained.

If it is asked, however, how that which in ourselves and in the remainder of the animal world is intellectual and perceptive, which thinks and wills, is ascribable to a mechanical process of organic development—whether the development of the mind can be conceived as resulting from purely mechanical laws? I answer unhesitatingly in the affirmative with the pure materialist, although I do not agree with him as to the manner in which he derives these phenomena from matter, since thinking and extension are heterogeneous things, and one cannot be considered as a product of the other. But why should not the ancient notion of “conscious matter” given out by Maupertuis and Robinet, not be again entertained, as pointed out in recent times by Fechner?[313] Would there not thus be found a useful formula for explaining phenomena hitherto quite incomprehensible?

Von Hartmann in criticizing himself, designates the sensibility of atoms as an “almost inevitable hypothesis” (p. 62), “inevitable because if sensibility were not a general and original property of the constituent elements of matter, it would be absolutely incomprehensible how through its potentiality and integration that sensibility known to us as being possessed by the organism could have arisen.” “It is impossible that from purely external elements devoid of all internality (Innerlichkeit) there should suddenly appear, by a certain mode of combination, an internality which becomes more and more richly developed. The more certainly science becomes convinced that in the sphere of externality (Äusserlichkeit) the higher (organic) phenomena are only results of combination, or are the aggregate phenomena of the elementary atomic forces, the more surely, when she once seriously concerns herself with this other question, will she not fail to be convinced that the sensibility possessed by higher stages of consciousness can be only combination-results, or the aggregate phenomena of the elementary sensations of atoms, although these atomic sensations as such always remain below the level of the higher combinations of consciousness.” In confusing this double-sided nature of the objective phenomenon “lies the main error of all materialism and of all subjective idealism. Just as the attempt of the latter (subjective idealism) to construct the external phenomena of existence in space out of functions of internality and their combinations is impossible, so is the endeavour of the former (materialism) to build up internal sensation out of any combinations of force acting externally in space equally impossible.”

I have no intention of going any deeper into these questions. I mention them only in order to point out that even from this side there appears to me no obstacle in the way of a purely mechanical conception of the processes of the universe. The naturalist may be excused if he attempts to penetrate into the region of philosophy; it arises from the wish to be able to contribute a little towards the reconciliation of the latest knowledge of the naturalist with the religious wants of the human mind—towards the aim striven for by both sides, viz. a satisfactory and harmonious view of the universe, according with the state of knowledge of our time.

I believe that I have shown that the theory of selection by no means leads—as is always assumed—to the denial of a teleological Universal Cause and to materialism, and I thereby hope that I have cleared the way for this doctrine, the importance of which it is scarcely possible to over-estimate. Many, and not the most ill-informed, do not get so far as to make an unbiassed examination into the facts, because they are at the outset alarmed by the to them inevitable consequence of the materialistic conception of the universe. Mechanism and teleology do not exclude one another, they are rather in mutual agreement. Without teleology there would be no mechanism, but only a confusion of crude forces; and without mechanism there would be no teleology, for how could the latter otherwise effect its purpose?[314]

Von Hartmann correctly says:—“The most complete mechanism conceivable is likewise the most completely conceivable teleology.” We may thus represent the phenomenal universe as such a completely conceivable mechanism. With this conception vanish all apprehensions that the new views would cause man to lose the best that he possesses—morality and purely human spiritual culture. He who, with Von Baer, considers the laws of nature as the “permanent expressions of the will of a creative principle,” will clearly perceive that a further advance in the knowledge of these laws need not divert man from the path of increasing improvement, but must further him in this course—that the knowledge of truth, whatever may be its purport, cannot possibly be considered a backward step. Let us take our stand boldly on the ground of new knowledge, and accept the direct consequences thereof, and we shall not be obliged to give up either morality or the comforting conviction of being part of an harmonious world, as a necessary member capable of development and perfection.

Any other mode of interference by a directive teleological power in the processes of the universe than by the appointment of the forces producing them, is however, at least to the naturalist, inadmissible. We are still far removed from completely understanding the mechanism by means of which the organic world is evoked—we still find ourselves at the very beginning of knowledge. We are, however, already convinced that both the organic and the inorganic worlds are dependent only upon mechanical forces, for to this conclusion we are led, not only by the results of investigators who have restricted themselves to limited provinces, but also by the most general considerations. But although the force of these arguments may not be acknowledged, and although one might maintain that the inductional proofs against the existence of a “phyletic vital force” have been directed only against points of detail, or have never been completely demonstrated, i.e. for all points, it must nevertheless be conceded, that for the naturalist the mechanical conception of Nature is the only one possible—that he is not at all justified in abandoning this view so long as the interference of teleological forces in the course of the processes of organic development has not been demonstrated to him. Thus, it will not be immaterial whether a conception of Nature which to many seems inevitable is consistent with the idea of universal design, or a final directive universal principle, since the value which we may attach to our own lives and aims, essentially depends thereon. The final and main result of this essay will thus be found in the attempted demonstration that the mechanical conception of Nature very well admits of being united with a teleological conception of the Universe.

THE END.

[FOOTNOTES]

[1] A most minute and exact description of the newly hatched larva of Chionobas Aëllo is given by the American entomologist, Samuel H. Scudder. Ann. Soc. Ent. de Belgique, xvi., 1873.

[2] I am aware that this certainly cannot be said of philosophers like Lotze or Herbert Spencer; but these are at the same time both naturalists and philosophers.

[3] “Über die Artrechte des Polyommatus Amyntas und Polysperchon.” Stett. ent. Zeit. 1849. Vol. x. p. 177–182. [In Kirby’s “Synonymic Catalogue of Diurnal Lepidoptera” Plebeius Amyntas is given as a synonym and P. Polysperchon as a var. of P. Argiades Pall. R.M.]

[4] “Die Arten der Lepidopteren-Gattung Ino Leach, nebst einigen Vorbemerkungen über Localvarietäten.” Stett. ent. Zeit. 1862. Vol. xxiii. p. 342.

[5] [Eng. ed. W. H. Edwards has since pointed out several beautiful cases of seasonal dimorphism in America. Thus Plebeius Pseudargiolus is the summer form of P. Violacea, and Phyciodes Tharos the summer form of P. Marcia. See Edwards’ “Butterflies of North America,” 1868–79.]

[6] [Eng. ed. I learn by a written communication from Dr. Speyer that two Geometræ, Selenia Tetralunaria and S. Illunaria Hüb., are seasonally dimorphic. In both species the winter form is much larger and darker.] [Selenia Lunaria, S. Illustraria, and some species of Ephyra (E. Punctaria and E. Omicronaria) are likewise seasonally dimorphic. For remarks on the case of S. Illustraria see Dr. Knaggs in Ent. Mo. Mag., vol. iii. p. 238, and p. 256. Some observations on E. Punctaria were communicated to the Entomological Society of London by Professor Westwood in 1877, on the authority of Mr. B. G. Cole. See Proc. Ent. Soc. 1877, pp. vi, vii. R.M.]

[7] [In 1860 Andrew Murray directed attention to the disguising colours of species which, like the Alpine hare, stoat, and ptarmigan, undergo seasonal variation of colour. See a paper “On the Disguises of Nature, being an inquiry into the laws which regulate external form and colour in plants and animals.” Edinb. New Phil. Journ., Jan. 1860. In 1873 I attempted to show that these and other cases of “variable protective colouring” could be fairly attributed to natural selection. See Proc. Zoo. Soc., Feb. 4th, 1873, pp. 153–162. R.M.]

[8] [A phenomenon somewhat analogous to seasonal change of protecting colour does occur in some Lepidoptera, only the change, instead of occurring in the same individual, is displayed by the successive individuals of the same brood. See Dr. Wallace on Bombyx Cynthia, Trans. Ent. Soc. Vol. v. p. 485. R.M.]

[9] “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872, pp. 55–62.

[10] [Mr. A. R. Wallace maintains that the obscurely coloured females of those butterflies which possess brightly coloured males have been rendered inconspicuous by natural selection, owing to the greater need of protection by the former sex. See “Contributions to the Theory of Natural Selection,” London, 1870, pp. 112–114. It is now generally admitted that the underside of butterflies has undergone protectional adaptation; and many cases of local variation in the colour of the underside of the wings, in accordance with the nature of the soil, &c., are known. See, for instance, Mr. D. G. Rutherford on the colour-varieties of Aterica Meleagris (Proc. Ent. Soc. 1878, p. xlii.), and Mr. J. Jenner Weir on a similar phenomenon in Hipparchia Semele (loc. cit. p. xlix.) R.M.]

[11] [The fact that moths which, like the Geometræ, rest by day with the wings spread out, are protectively marked on the upper side, fully corroborates this statement. R.M.]

[12] “Über die Einwirkung verschiedener, während der Entwicklungsperioden angewendeter Wärmegrade auf die Färbung und Zeichnung der Schmetterlinge.” A communication to the Society of Natural Science of Steiermark, 1864.

[13] See Exp. 9, [Appendix I].

[14] See Exp. 11, [Appendix I].

[15] See Exps. 4, 9, and 11, [Appendix I].

[16] It seems to me very necessary to have a word expressing whether a species produces one, two, or more generations in the year, and I have therefore coined the expression mono-, di-, and polygoneutic from γονεύω, I produce.

[17] [Eng. ed. In the German edition, which appeared in 1874, I was not able to support this hypothesis by geographical data, and could then only ask the question “whether in the most northern portion of its area of distribution, appears in two or only in one generation?” This question is now answered by the Swedish Expedition to the Yenisei in 1876. Herr Philipp Trybom, one of the members of this expedition, observed A. Levana at the end of June and beginning of July, in the middle of Yenisei, in 60°-63° N. (Dagfjärilar från Yenisei in Översigt ap k. Vertensk. Akad. Förhandlingon, 1877, No. 6.) Trybom found Levana at Yenisk on June 23rd, at Worogova (61° 5´) on July 3rd, at Asinova (61° 25´) on July 4th, at Insarowa (62° 5´) on July 7th, and at Alinskaja (63° 25´) on July 9th. The butterflies were especially abundant at the beginning of June, and were all of the typical Levana form. Trybom expressly states, “we did not find a single specimen which differed perceptibly from Weismann’s [Figs. 1 and 2] (‘Saison-Dimorphismus’ Taf. I.).”

The Swedish expedition soon left the Yenisei, and consequently was not able to decide by observations whether a second generation possessing the Prorsa form appeared later in the summer. Nevertheless, it may be stated with great probability that this is not the case. The districts in which Levana occurs on the Yenisei have about the same isotherm as Archangel or Haparanda, and therefore the same summer temperature. Dr. Staudinger, whose views I solicited, writes to me:—“In Finnmark (about 67° N.) I observed no species with two generations; even Polyommatus Phlæas, which occurs there, and which in Germany has always two, and in the south, perhaps, three generations, in Finnmark has only one generation. A second generation would be impossible, and this would also be the case with Levana in the middle of Yenisei. I certainly have Levana and Prorsa from the middle of Amur, but Levana flies there at the end of May, and the summers are very warm.” The middle of Amur lies, moreover, in 50° N. lat., and therefore 10°-13° south of the districts of the Yenisei mentioned.

It must thus be certainly admitted that on the Yenisei A. Levana occurs only in the Levana form, and that consequently this species is at the present time, in the northernmost portion of its area of distribution, in the same condition as that in which I conceive it to have been in mid Europe during the glacial period. It would be of the greatest interest to make experiments in breeding with this single-brooded Levana from the Yenisei, i.e., to attempt to change its offspring into the Prorsa form by the action of a high temperature. If this could not be accomplished it would furnish a confirmation of my hypothesis than which nothing more rigorous could be desired.]

[18] See Exp. 10, [Appendix I].

[19] When Dorfmeister remarks that hibernating pupæ which, at an early stage “were taken for development into a room, or not exposed to any cold, gave dwarfed, weakly and crippled,” or otherwise damaged butterflies, this is entirely attributable to the fact that this able entomologist had neglected to supply the necessary moisture to the warm air. By keeping pupæ over water I have always obtained very fine butterflies.

[20] [For other remarkable cases of sexual dimorphism (not antigeny in the sense used by Mr. S. H. Scudder, Proc. Amer. Acad., vol. xii. 1877, pp. 150–158) see Wallace “On the Phenomena of Variation and Geographical Distribution, as illustrated by the Papilionidæ of the Malayan Region,” Trans. Linn. Soc., vol. xxv. 1865, pp. 5–10. R.M.]

[21] [Eng. ed. Dimorphism of this kind has since been made known: the North American Limenitis Artemis and L. Proserpina are not two species, as was formerly believed, but only one. Edwards bred both forms from eggs of Proserpina. Both are single-brooded, and both have males and females. The two forms fly together, but L. Artemis is much more widely distributed, and more abundant than L. Proserpina. See “Butterflies of North America,” vol. ii.]

[22] [Eng. ed. Edwards has since proved experimentally that by the application of ice a large proportion of the pupæ do indeed give rise to the var. Telamonides. He bred from eggs of Telamonides 122 pupæ, which, under natural conditions, would nearly all have given the var. Marcellus. After two months’ exposure to the low temperature there emerged from August 24th to October 16th, fifty butterflies, viz. twenty-two Telamonides, one intermediate form between Telamonides and Walshii, eight intermediate forms between Telamonides and Marcellus more nearly related to the former, six intermediate forms between Telamonides and Marcellus, but more closely resembling the latter, and thirteen Marcellus. Through various mishaps the action of the ice was not complete and equal. See the “Canadian Entomologist,” 1875, p. 228. In the newly discovered case of Phyciodes Tharos also, Edwards has succeeded in causing the brood from the winter form to revert, by the application of ice to this same form. See [Appendix II]. for a résumé of Edwards’ experiments upon both Papilio Ajax and Phyciodes Tharos. R.M.]

[23] Thus from eggs of Walshii, laid on April 10th, Edwards obtained, after a pupal period of fourteen days, from the 1st to the 6th of June, fifty-eight butterflies of the form Marcellus, one of Walshii, and one of Telamonides.

[24] [The word ‘Amixie,’ from the Greek ἀμιξία, was first adopted by the author to express the idea of the prevention of crossing by isolation in his essay “Über den Einfluss der Isolirung auf die Artbildung,” Leipzig, 1872, p. 49. R.M.]

[25] [Eng. ed. In 1844, Boisduval maintained this relationship of the two forms. See Speyer’s “Geographische Verbreit. d. Schmetterl.,” i. p. 455.]

[26] According to a written communication from Dr. Staudinger, the female Bryoniæ from Lapland are never so dusky as is commonly the case in the Alps, but they often have, on the other hand, a yellow instead of a white ground-colour. In the Alps, yellow specimens are not uncommon, and in the Jura are even the rule.

[27] [According to W. F. Kirby (Syn. Cat. Diurn. Lepidop.), the species is almost cosmopolitan, occurring, as well as throughout Europe, in Northern India (var. Timeus), Shanghai (var. Chinensis), Abyssinia (var. Pseudophlæas), Massachusetts (var. Americana), and California (var. Hypophlæas). In a long series from Northern India, in my own collection, all the specimens are extremely dark, the males being almost black. R.M.]

[28] [Eng. ed. From a written communication from Dr. Speyer, it appears that also in Germany there is a small difference between the two generations. The German summer brood has likewise more black on the upper side, although seldom so much as the South European summer brood.]

[29] [Assuming that in all butterflies similar colours are produced by the same chemical compounds. R.M.]

[30] [Mr. H. W. Bates mentions instances of local variation in colour affecting many distinct species in the same district in his memoir “On the Lepidoptera of the Amazon Valley;” Trans. Linn. Soc., vol. xxiii. Mr. A. R. Wallace also has brought together a large number of cases of variation in colour according to distribution, in his address to the biological section of the British Association at Glasgow in 1876. See “Brit. Assoc. Report,” 1876, pp. 100–110. For observations on the change of colour in British Lepidoptera according to distribution see papers by Mr. E. Birchall in “Ent. Mo. Mag.,” Nov., 1876, and by Dr. F. Buchanan White, “Ent. Mo. Mag.,” Dec., 1876. The colour variations in all these cases are of course not protective as in the well-known case of Gnophos obscurata, &c. R.M.]

[31] See Figs. 10 and 14, 11 and 15, [Plate I].

[32] “On the Origin and Metamorphoses of Insects,” London, 1874.

[33] I at first thought of designating the two forms of cyclical or homochronic heredity as ontogenetic- and phyletic-cyclical heredity. The former would certainly be correct; the latter would be also applicable to alternation of generation (in which actually two or more phyletic stages alternate with each other) but not to all those cases which I attribute to heterogenesis, in which, as with seasonal dimorphism, a series of generations of the same phyletic stage constitute the point of departure.

[34] When Meyer-Dürr, who is otherwise very accurate, states in his “Verzeichniss der Schmetterlinge der Schweiz,” (1852, p. 207), that the winter and summer generations of P. Ægeria differ to a small extent in the contour of the wings and in marking, he has committed an error. The characters which this author attributes to the summer form are much more applicable to the female sex. There exists in this species a trifling sexual dimorphism, but no seasonal dimorphism.

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

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

[37] [Trans. Linn. Soc., vol. xxv. 1865, p. 9. R.M.]

[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.”

[39] Loc. cit. chap. iv.

[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.

[41] See my essay “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872.

[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.]

[43] “Grundzüge der Zoologie.” 2nd ed. Leipzig, 1872. Introduction.

[44] With reference to this subject, see the discussion by the Belgian Entomological Society, Brussels, 1873.

[45] P. E. Müller, “Bidrag til Cladocerners Fortplantingshistorie,” 1868.

[46] Sars, in “Förhandlinger i Videnskabs Selskabet i Christiania,” 1873, part i.

[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.]

[48] See my memoir, “Über Bau und Lebenserscheinungen der Leptodora hyalina,” Zeitschrift f. wiss. Zool., vol. xxiv. part 3, 1874.

[49] Stettin. entom. Zeit., vol. xviii. p. 83, 1857.

[50] Compt. Rend., vol. lxxvii. p. 1164, 1873.

[51] [“Accidental” in the sense of our being in ignorance of the laws of variation, as so frequently insisted upon by Darwin. R.M.]

[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.]

[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.

[54] See my essay, “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872.

[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.]

[56] See my essay, “Über die Berechtigung der Darwin’schen Theorie.” Leipzig, 1868.

[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.

[58] The experiments upon Papilio Ajax and Phyciodes Tharos, described in this Appendix, were made by Mr. W. H. Edwards (see his “Butterflies of North America;” also the “Canadian Entomologist,” vol. vii. p. 228–240, and vol. ix. p. 1–10, 51–5, and 203–6); and I have added them, together with some hitherto unpublished results, to Dr. Weismann’s Essay, in order to complete the history of the subject of seasonal dimorphism up to the present time.—R.M.

[59] This is a striking illustration of the diversity of individual constitution so frequently insisted on by Dr. Weismann in the foregoing portion of this work.

[60] The reader who wishes to acquire a detailed knowledge of the different varieties of this butterfly, of which a very large number are known, must consult the plates and descriptions in Edwards’ “Butterflies of North America,” vol. ii.

[61] Mr. Edwards has shown also that Argynnis Myrina can lay fertile eggs when but a few hours out of the chrysalis. Canad. Ent., September, 1876, vol. viii. No. 9.

[62] Mr. Edwards remarks that the habit of becoming lethargic is of great service to a digoneutic species in a mountain region where it is exposed to sharp changes of temperature. “If the fate of the species depended on the last larval brood of the year, and especially if the larvæ must reach a certain stage of growth before they were fitted to enter upon their hibernation, it might well happen that now and then an early frost or a tempestuous season would destroy all the larvæ of the district.”

[63] Compare this with Weismann’s remarks, pp. [19]–[22], and [53].

[64] See Canad. Ent., vol. ix. p. 69.

[65] Figures of the different forms of this species are given in vol. i. of Edward’s “Butterflies of North America.”

[66] Only the species of Smerinthus can be made to lay eggs regularly in confinement; Macroglossa Stellatarum laid a number in a large gauze-covered breeding-cage; the species of Deilephila could not be induced to lay more than single ones in such a cage. From species of Chærocampa also I never obtained but a few eggs, and from Sphinx and Acherontia never more than single ones.

[67] [Eng. ed. Since the appearance of the German edition of this work, numerous descriptions of the young stages of caterpillars have been given, but in all cases without representing the relationship of the forms.] [In the excellent figures of larvæ at various stages of growth, given in some of the more recent works on Lepidoptera, there will be found much material which may be regarded as a contribution to the field of research entered on by the author in the present essay, i.e. the ontogeny and comparative morphology of larval markings, although it is much to be regretted that the figures and descriptions have not been given from this point of view. In his “Butterflies of North America,” for example, W. H. Edwards figures the young as well as the adult larvæ of species of Apatura, Argynnis, Libythea, Phyciodes, Limenitis, Colias, Papilio, &c. Burmeister, in his recently published “Lépidoptères de la République Argentine,” figures the young stages of species of Caligo, Opsiphanes, Callidryas, Philampelus, &c. Messrs. Hellins and Buckler have figured and described the early stages of large numbers of the caterpillars of British Lepidoptera, but their figures remain unpublished. The larvæ of many of our native species belonging to the genera Liparis, Tæniocampa, Epunda, Cymatophora, Calocampa, &c., are dull when young, but become brightly coloured at the last moult. Such changes of colour are probably associated with some change, either in the habits or in the environment; and a careful study of the ontogenetic development of such species in connection with their life-history would furnish results of great value to the present inquiry. The same remarks apply to those Noctuæ larvæ which are brightly coloured in their young stages, and become dull when adult.

Among other papers which may be considered as contributions to the present subject, I may mention the following:—In 1864 Capt. Hutton published a paper, “On the Reversion and Restoration of the Silkworm, Part II.” (Trans. Ent. Soc. 1864, p. 295), in which he describes the various stages of development of several species of Bombycidæ. In 1867 G. Semper published accounts of the early stages of several Sphinx-larvæ (“Beiträge zur Entwicklungsgeschichte einiger ostasiatischer Schmetterlinge,” Verhandl. k.k. Zoolog.-botan. Gesell. in Wien, vol. xvii.). The question as to the number of claspers in young Noctuæ larvæ has been raised in notes by Dr. F. Buchanan White (“Ent. Mo. Mag.,” vol. v. p. 204) and B. Lockyer (“Entomologist,” 1871, p. 433). A valuable paper, “On the Embryonic Larvæ of Butterflies,” was published in 1871 by S. H. Scudder (“Ent. Mo. Mag.,” vol. viii. p. 122). For remarks on the development of the larva of Papilio Merope, see J. P. Mansel Weale in Trans. Ent. Soc., 1874, p. 131, and Pl. I.; also this author on the young stages of the larva of Gynanisa Isis, Trans. Ent. Soc., 1878, p. 184. For an account of the development of the larvæ of certain North American species of Satyrus, see W. H. Edwards in the “Canadian Entom.,” vol. xii. p. 21. Mr. P. H. Gosse’s recent description of the newly hatched caterpillar of Papilio Homerus (Proc. Ent. Soc. 1879, p. lv), furnishes a good illustration of the value of studying the ontogeny. The natural affinities of the Papilionidæ were at one time much disputed, some systematists placing this family at the head of the Lepidoptera, and others regarding them as being more closely allied to the moths. Mr. Gosse’s observation tends to confirm the latter view, now generally received by Lepidopterists, since he states that the larva in question “suggests one of the great Saturniadæ, such as Samia Cecropia.” Mr. Scudder, in the paper above referred to, adopts an analogous argument to show the close relationship between the Papilionidæ and Hesperidæ. R.M.]

[68] [Mr. A. G. Butler has recently furnished a good illustration of the danger of classifying Lepidoptera according to the affinities of the perfect insects only, in his paper, “On the Natural Affinities of the Lepidoptera hitherto referred to the Genus Acronycta of authors,” Trans. Ent. Soc. 1879, p. 313. If the author’s views are ultimately accepted, the species at present grouped under this genus will be distributed among the Arctiidæ, Liparidæ, Notodontidæ, and Noctuæ. Mr. Butler’s determination of the affinities of the species supposed to belong to the genus mentioned, is based chiefly upon a comparative examination of the larvæ, and this is far more likely to show the true blood-relationship of the species than a comparison of the perfect insects only. A study of the comparative ontogeny can alone give a final answer to this question. R.M.]

[69] [In his recent revision of the Sphingidæ, Mr. A. G. Butler (Trans. Zoo. Soc., vol. ix. part x.) retains Walker’s arrangement. R.M.]

[70] The deposition of black pigment may commence immediately before ecdysis.

[71] [Mr. Herbert Goss states (Proc. Ent. Soc. 1878, p. v.) that according to his experience, the green and brown varieties of C. Porcellus (erroneously printed as Elpenor in the passage referred to) are about equally common, the former colour not being in any way confined to young larvæ. Mr. Owen Wilson in his recent work, “The Larvæ of British Lepidoptera and their food-plants,” figures ([Pl. VIII]., Figs. 3 and 3a) the two forms, both apparently in the adult state. During the years 1878–79, my friend, Mr. J. Evershed, jun., took five of these full-grown larvæ in Surrey, one of these being the green variety. In order to get more statistics on this subject, I applied this year (1880) to Messrs. Davis of Dartford, who informed me that among 18–20 adult caterpillars of Porcellus in their possession, there was only one green specimen. R.M.]

[72] I unite the genera Pergesa and Darapsa of Walk. with Chærocampa, Dup.; the first appears to me to be quite untenable, since it is impossible that two species, of which the caterpillars agree so completely as those of C. Elpenor and Porcellus, can be located in different genera. Porcellus indeed was referred to the genus Pergesa because of its different contour of wings, an instance which distinctly shows how dangerous it is to attempt to found Lepidopterous genera without considering the caterpillars. The genus Darapsa also appears to me to be of very doubtful value, and in any case requires further confirmation with respect to the larval forms.

[73] [Mr. A. G. Butler (Trans. Zoo. Soc., vol. ix., part. x., 1876) gives a list of about eighty-four species of Chærocampa, and sixteen of Pergesa, besides numerous other species belonging to several genera placed between Chærocampa and Pergesa. Of Darapsa, he states “that this genus was founded upon most heterogeneous material, the first three species being referable to Hübner’s genus Otus, the fifth to Walker’s genus Diodosida, the sixth and eighth to the genus Daphnis of Hübner, the seventh, ninth, and tenth to Chærocampa of Duponchel; there therefore remains only the fourth species, allied to Chærocampa, but apparently sufficiently distinct.” The species still retained in the genus Darapsa is D. rhodocera, Wlk., from Haiti. R.M.]

[74] [Otus Syriacus of Butler’s revision. R.M.]

[75] Abbot and Smith. “The Natural History of the rarer Lepidopterous Insects of Georgia, collected from the observations of John Abbot, with the plants on which they feed.” London, 1797, 2 vols. fol.

[76] [Otus Chœrilus and O. Myron of Butler’s revision. R.M.]

[77] [To this group may also be added Ampelophaga Rubiginosa, Ménétriés, from China and Japan, the caterpillar of which, having the distinct subdorsal line without any trace of eye-spots, is figured by Butler (loc. cit., Pl. XCI., Fig. 4). This author also gives a figure of another species belonging to the subfamily Chærocampinæ (Pl. XC., Fig. 11), viz. Acosmeryx Anceus, Cram., from Amboina, Java, Silhet, and S. India; the caterpillar is green, with seven oblique yellow stripes along the sides, and a very conspicuous white subdorsal line with a red border above. As there are no eye-spots, this species may be referred to the present group provisionally, although its general marking is very distinct from that of the Chærocampa group. R.M.]

[78] [Eng. ed. Dr. Staudinger has since obtained the caterpillar of C. Alecto from Beyrout; it possesses “a very distinct subdorsal line, and on the fourth segment a beautiful eye-spot, which is repeated with gradual diminution to segments 7–8”.]

[79] Figured in “A Catalogue of Lepidopterous Insects in the Museum of the East India Company,” by Thomas Horsfield and Frederick Moore. London, 1857. Vol. i., Pl. XI.

[80] Figured in Trans. Ent. Soc., New Series, vol. iv., Pl. XIII.

[81] Ibid.

[82] [The following species figured by Butler (loc. cit. Pls. XC. and XCI.) appear to belong to the second group—Chærocampa Japonica, Boisd., which is figured in two forms, one brown, and the other green. The former has two distinct ocelli on the fourth and fifth segments, and a distinct rudiment on the sixth, whilst the subdorsal line extends from the second eye-spot to the caudal horn, and beneath this line the oblique lateral stripes stand out conspicuously in dark brown on a lighter ground. The ocelli are equally well developed on the fourth and fifth segments in the green variety, the subdorsal line commencing on the sixth segment, and extending to the caudal horn; there is no trace of a third eye-spot, nor are there any oblique lateral stripes; the insect is almost the exact counterpart of C. Elpenor in its fourth stage. (See Fig. 21, [Pl. IV].) Pergesa Mongoliana, Butl., is brown, without a trace of the subdorsal line except on the three front segments, and with only one large eye-spot on the fourth segment. Chærocampa Lewisii, Butl., from Japan, is likewise figured in two forms. The brown variety has the subdorsal line on the three front segments only, distinct ocelli on the fourth and fifth segments, and gradually diminishing rudiments on the remaining segments. The green form appears to be transitional between the present and the third group, as it possesses a distinct, but rudimentary eye-spot on the third segment, besides the fully developed ones on the fourth and fifth, and very conspicuous, but gradually decreasing repetitions of rudimentary ocelli on segments 6–10. To this group may be added Chærocampa Aristor, Boisd., the caterpillar of which is figured by Burmeister (Lép. Rép. Arg., Pl. XV., Fig. 4) in the characteristic attitude of alarm, with the front segments retracted, and the ocelli on the fourth segment prominently exposed. The subdorsal line is present in this species. Burmeister also figures two of the early stages (Pl. XV., Fig. 7, A and B), and describes the complete development of Philampelus Labruscæ, another species belonging to the subfamily Chærocampinæ. The earliest stage (3–4 days old) is simple green, with no trace of any marking except a black spot on each side of the fourth segment, the position of the future ocelli. A curved horn is present both in this stage and the following one, during which the caterpillar is still green, but now has seven oblique red lateral stripes. The caudal horn is shed at the second moult, after which the colour becomes darker, the adult larva (figured by Madame Mérian, in her work on Surinam, pl. 34 and Sepp., pl. 32) being mottled brown. In addition to the ocellus on the fourth segment, there is another slightly larger on the eleventh segment, so that this species may perhaps be another transition to the third group; but our knowledge is still too imperfect to attempt to generalize with safety. R.M.]

[83] Cat. Lep. Ins. East Ind. Comp., Pl. XIII. [Figured also by Butler (=Chæerocampa Silhetensis, Walker), loc. cit. Pl. XCII., Fig. 8. R.M.]

[84] Cat. Lep. Ins. East Ind. Comp., Pl. XIII. [Figured also by Butler, loc. cit. Pl. XCI., Fig. 1. R.M.]

[85] Horsfield and Moore, loc. cit. Pl. X.

[86] Ibid. [=Pergesa Acteus, Walker. R.M.]

[87] [Figured also by Burmeister, loc. cit. Pl. XV., Fig. 3. R.M.]

[88] Horsfield and Moore, loc. cit., Pl. XI.

[89] To be accurate this should be designated the infra-spiracular line; but this term cannot be well applied except in cases where there is also a supra-spiracular line, as, for instance, in Anceryx (Hyloicus) Pinastri.

[90] Upon this fact obviously depends the statement of that extremely accurate observer Rösel, that the caterpillar of Euphorbiæ is but very slightly variable (“Insektenbelustigungen,” Bd. iii. p. 36). I formerly held the same opinion, till I convinced myself that this species is very constant in some localities, but very variable in others. It appears that local influences make the caterpillar variable.

[91] The green is considerably too light in [Fig. 45].

[92] “Die Pflanzen und Raupen Deutschlands.” Berlin, 1860, p. 83.

[93] Fig. 62, [Pl. VII]., is copied from Boisduval.

[94] The fading of the red anteriorly has not been represented in the figure.

[95] [The caterpillar of Deilephila Euphorbiarum, figured by Burmeister (Lép. Rép. Arg., Pl. XVI, Fig. 1) belongs to this stage. R.M.]

[96] [In concluding this account of the Chærocampinæ I may call attention to the following species, which have since been figured by Burmeister:—Pachylia Ficus, Linn. (loc. cit. Pl. XIV., Figs. 1 and 2); during the three first stages the larva is uniformly green, with a yellow subdorsal line, and below this ten oblique yellow stripes slanting away from the head; after the third moult the colour completely changes, the whole area of the body being divided into two distinct portions by the subdorsal line, above which the colour is red, and underneath of a pale green; the oblique stripes have almost disappeared; no occelli nor annuli are present. Pachylia Syces, Hübn. (loc. cit. Fig. 3); very similar to the last species in its young stages (figured also by Mérian, Surin. pl. 33). Philampelus Vitis, Linn. (loc. cit. Figs. 4 and 5); two stages represented; between first and second moults green, with oblique paler stripes slanting in same direction as in Pachylia, and each one containing a red streak surrounding the spiracle. When adult, the ground-colour is yellow above and green beneath, the whole surface being mottled with deep black and red transverse markings; the oblique stripes whitish, bordered with black at their lower extremities (figured also by Mérian, pls. 9 and 39). Philampelus Anchemolus, Cram. (loc. cit. Pl. XV., Fig. 1; Mérian, pl. 47); green when young, with seven oblique red stripes; when adult, uniformly brown, with seven pale yellow lateral markings, the first four of which are spots, and the remainder broad oblique stripes slanting forwards. Philampelus Labruscæ, (see note [82], p. [195]). R.M.]

[97] [Mimas Tiliæ of Butler’s revision. The author states that this genus is “easily distinguished from Laothoë by the form of the wings, the outer margin of secondaries deeply excavated below the apex, and the secondaries narrow and not denticulated.” Here again we have a clashing of the results arrived at by a study of the ontogeny of the larvæ, on the one hand, and the founding of genera on the characters of the imagines only, on the other. Of the three species discussed by Dr. Weismann, Mr. Butler, following other authors, refers Tiliæ to the genus Mimas, Populi to Laothoë, and Ocellatus to Smerinthus. It is to be hoped that when our knowledge of the developmental history of larvæ is more complete in all groups, a reconciliation between the results of the biological investigator and the pure systematist will be brought about, so that a genus may not, as at present, have such very different values when regarded from these two points of view. R.M.]

[98] The caterpillar is thus figured by Rösel.

[99] [In 1879 Mr. E. Boscher found about thirty full-grown caterpillars of this species in the neighbourhood of Twickenham, ten to twelve of which were feeding on Salix viminalis, and the remainder, from a locality not far distant, on Salix triandra. The whole of the specimens taken on the plant first named, had the red-brown spots above and below the oblique stripes more or less completely developed, as I myself had an opportunity of observing. In these spotted specimens the ground-colour was bright yellowish-green, and in the others this colour was dull whitish-green above, passing into bluish-green below. Should these observations receive wider confirmation, it would be fair to conclude that this species is now in two states of phyletic development, the more advanced stage being represented by the brighter spotted variety. (See also Proc. Ent. Soc. 1879, p. xliv.). Mr. Peter Cameron has recently suggested (Trans. Ent. Soc. 1880, p. 69) that the reddish-brown spots on the Smerinthus caterpillars may serve for purposes of disguise, as they closely resemble, both in colour and form, certain galls (Phytoptus) of the food-plants of these species. If this view be admitted, these spots must be considered as a new character, now being developed by natural selection. The variation in the ground-colour of the two forms of S. Ocellatus may possibly be phytophagic, but this can only be decisively settled by a series of carefully conducted experiments. R.M.]

[100] “Insekten-Belustigungen,” Suppl. Pl. 38, Fig. 40.

[101] “Catalogue of Lepidop.” British Museum. [Butler divides the subfamily Smerinthinæ into 17 genera, containing 79 species, viz. Metamimas, 2; Mimas, 4; Polyptychus, 7; Lophostethus, 1; Sphingonæpiopsis, 1; Langia, 2; Triptogon, 23; Laothoë, 2; Cressonia, 3; Paonias, 2; Calasymbolus, 5; Smerinthus, 5; Pseudosmerinthus, 2; Daphnusa, 4; Leucophlebia, 5; Basiana, 10; Cæquosa, 1. R.M.]

[102] “Cabinet Orient. Entom.,” p. 13, [Pl. VI]., Fig. 2. [Butler places this species doubtfully among the Sphinginæ. R.M.]

[103] “Catalogue of the Lepidop. Insects of the E.I. Co.,” by Horsfield and Moore. [Pl. VIII]., Fig. 6.

[104] [The larvæ of four other species of this subfamily have since been made known through Mr. Butler’s figures. Smerinthus Tatarinovii, Ménetriés (loc. cit. Pl. XC., Fig. 16), from Japan, is “pale sea-green, tuberculated with white, with seven lateral, oblique, crimson-edged white stripes.” There is no trace of the subdorsal line shown in the figure, so that this species thus appears to be in the third phyletic stage of development. Smerinthus Planus, Walker, from China (loc. cit. Pl XCII., Fig. 11), is “pale green, with white or yellow lateral stripes.” A trace of the subdorsal line remains on the front segments, thus showing that the species is in the second phyletic stage of development. Triptogon Roseipennis, Butler, from Hakodadi (loc. cit. Pl. XCI., Fig. 6), is represented as yellow, with seven oblique white stripes, with large irregular triangular red spots extending from the anterior edge of the stripes, nearly across each segment. It is probably in the third phyletic stage. The Indian Polyptychus Dentatus, Cramer (loc. cit. Pl. XCI., Fig. 10), is “bluish-green at the sides, with oblique purple stripes, with a broad, dorsal, longitudinal, golden-green band, bordered by subtriangular purple spots, one above each stripe.” The dorsal band is bordered by coloured stripes, which may be the subdorsal lines; but the position in which it is figured, and its very different mode of coloration, make it very difficult to compare satisfactorily with the foregoing species. The genus Ambulyx is closely allied to the Smerinthinæ, and the two following species may be here mentioned: A. Gannascus, Stoll, figured by Burmeister (loc. cit. Pl. XIII., Fig. 5), is green, with a yellow subdorsal line, and seven oblique white lateral stripes, edged with red. A. Liturata, Butl. (loc. cit. Pl. XCI., Fig. 2), is yellowish-green above, passing into bluish-green below. The subdorsal is present on the three front segments, and is followed by a row of white, elongated patches, one on each segment, these being the upper portions of a row of lateral oblique stripes. The thickened upper extremities of the latter are edged with red, and their arrangement is very suggestive of their having arisen from the breaking up of a subdorsal line. R.M.]

[105] [Butler catalogues 43 species of this genus. R.M.]

[106] The deposition of eggs was accomplished by the insect laying hold of the point of a twig with its legs during flight, and curving its abdomen upwards against a leaf, the wings being kept vibrating. The egg is instantaneously fastened to the leaf. This operation is repeated from twice to four times successively, the moth then hovering over and sucking at the flowers for some time. The eggs exactly resemble in colour the young green buds of Galium.

[107] [Figures of a remarkable case of gynandromorphism in a butterfly (Cirrochroa Aoris, Doubl.) have recently been published by Prof. Westwood (Trans. Ent. Soc. 1880, p. 113). On the right fore- and hind-wings of a male specimen there are patches of female colouring, thus bearing out in a very striking manner the above views concerning the non-fusibility of characters (in this case sexual) which have been long fixed. Complete (i.e. half-and-half) gynandromorphism is not uncommon in butterflies. R.M.]

[108] [I have long held the opinion that the di- and trimorphism displayed by certain butterflies has originated through polymorphism from ordinary variability. I will not here enter into details, but will only cite a few instances indicating the general direction of the arguments. The phenomenon to which I refer is that so ably treated of by Mr. A. R. Wallace (see Part I., p. [32], note [20]) and others. One male has often two or more distinctly coloured females, and in such cases one form of the female generally resembles the male in colour. Cases of polymorphic mimetic females may for the present be excluded, in order to reduce the argument to its greatest simplicity. Thus, in the case of native species, Colias Edusa has two females, one having the orange ground-colour of the male, and the other the well-known light form, var. Helice. So, also, Argynnis Paphia has a normal female and the dark melanic form var. Valezina. Numerous other cases might be mentioned among exotic species; and, looking at the phenomenon as a whole, it is seen to be one of gradation. For instance, our common “Blues” (Plebeius Icarus, P. Thetis, &c.) have females showing a complete gradation between the ordinary blue male and the brown female coloration. In a large number of specimens of Callosune Eupompe in my cabinet, collected in Arabia by the late J. K. Lord, there is a completely graduated series of females, varying from individuals having the scarlet tips of the fore-wings as strongly developed as in the males, to specimens without a trace of such colouring; and the same is the case with other species of this and allied genera. In such instances it is only necessary for the intermediate female forms to become extinct, in order to have true cases of dimorphism. It is significant that in 1877, when Colias Edusa appeared in this country in such extraordinary profusion, large numbers of intermediate forms were captured, these forming an uninterrupted series connecting the normal female and the var. Helice. R.M.]

[109] [Many of our best describers of caterpillars, such as the late Edward Newman, Messrs. Hellins and Buckler, &c., have described the various forms of numerous polymorphic species, but not from the point of view of the comparative morphology and ontogeny of the markings. R.M.]

[110] [In Butler’s revision both these species are placed in the genus Hemaris. R.M.]

[111] [This species is figured also by Butler (loc. cit. Pl. XC., Fig. 9), who represents it with seven oblique green lines between the spiracles and below the subdorsal line. R.M.]

[112] “Cat. E. Ind. Co. Mus.,” [Pl. VIII]., Fig. 2. [Walker, Lepidop. Heter. VIII., p. 92, No. 14, 1856; this species is strictly confined to Java. R.M.]

[113] [Eng. ed. The caterpillar is described and figured by Millière, “Iconographie des Chenilles et Lépidoptères inédits,” tome iii., Paris, 1869; also in the Annales, Soc. Linn. de Lyon, 1871 and 1873.] [This sp. = Hemaris Croatica, Esper., of Butler’s revision. R.M.]

[114] [The following additional species of the subfamily Macroglossinæ have been figured by Butler:—Lophura Hyas, Walk. (loc. cit. Pl. XC., Figs. 1 and 2), Hong-Kong, Silhet, and Java. The larva is apparently figured in two stages, the younger being red-brown with oblique white stripes, and the head and three front segments green. The larger specimen is green, mottled with red-brown, and no oblique stripes. In both figures the subdorsal line is indicated. The whole colouring is very suggestive of protective resemblance. Hemaris Hylas, Linn., from China, Japan, Ceylon, India, Australia, and Africa (loc. cit. Pl. XC., Fig. 4). The upper part of the body is light blue, and the lower part green, the two areas being separated by a white subdorsal line bordered above with brown. The dorsal line is feebly represented. Macroglossa Belis, Cram., N. India (loc. cit. Pl. XC., Fig. 6), is figured with the ground-colour deep indigo; a conspicuous white subdorsal, and a yellow spiracular line is present; on the side of each segment, between the two lines mentioned, there is a large red spot with a yellow nucleus (? eye-spots), the spots decreasing in size towards the head and tail; these probably confer upon this species some special protective advantage. Macroglossa Pyrrhosticta, Butler, China and Japan (loc. cit. Pl. XC., Fig. 8), is greenish-white with dorsal and subdorsal lines, and seven dark oblique stripes along the sides, below the subdorsal line. Of the foregoing species Hemaris Hyas appears to be in the same phyletic stage as M. Stellatarum and M. Croatica, &c., whilst M. Pyrrhosticta is probably, together with M. Corythus and M. Gilia, in another and more advanced stage, which is also passed through by Lophura Hyas in the course of its ontogenetic development. This last species (adult) and M. Belis may represent phyletic stages still further advanced. Caliomma Pluto, Walk., of which the caterpillar is figured by Burmeister (loc. cit. Pl. XIII., Fig. 1), appears to be a case of special protective resemblance to a twig or branch of its food-plant. Figured also by Chavannes; Bull. Soc. Vadoise des Sci. Nat., Dec. 6th 1854. R.M.]

[115] [Genus Pterogon, Boisd., = Proserpinus and Lophura (part). Butler, loc. cit. p. 632. The species above treated of = Proserpinus Œnotheræ, Fabr. R.M.]

[116] [These species = Thyreus Abboti and Proserpinus Gauræ of Butler’s revision. Of the former he states:—“Transformations described, and larva and imago figured, Am. Ent. ii. p. 123, 1870; the larva is also figured by Scudder in Harris’s ‘Correspondence,’ [Pl. III]., Fig. 1 (1869), and by Packard in his ‘Guide,’ p. 276, Fig. 203.” R.M.]

[117] [Proserpinus (Sphinx) Gorgon, Esp. R.M.]

[118] Rösel, loc. cit. vol. iii., p. 26, note.

[119] Figured and described by Abbot and Smith. [Macrosila (Sphinx) Cingulata is figured also by Burmeister, loc. cit. Pl. XII., Fig. 1. R.M.]

[120] Figured in “Cat. Lep. E. Ind. Co.”

[121] See the figure in Sepp’s Surinam Lepidoptera, P. 3, Pl. CI., 1848. A specimen in alcohol of the adult caterpillar is in the Berlin Museum. [The following is the synonymy of the above mentioned species:—Macrosila Hasdrubal, Walk. = Pseudosphinx (Sphinx) Tetrio, Linn.; M. Cingulata = Protoparce (Sphinx) Cingulata, Fabr.; M. Rustica = Protoparce (Sphinx) Rustica, Fabr.; Sphinx Convolvuli, Linn. = Protoparce Convolvuli; S. Carolina, Linn. = P. Carolina; the other species remain in the genera, as given above. The following additional species of Sphinginæ and Acherontiinæ have been figured by Butler:—Pseudosphinx Cyrtolophia, Butl., from Madras (loc. cit. Pl. XCI., Figs. 11 and 13); Protoparce Orientalis, Butl., from India, China, Java, &c. (Pl. XCI., Fig. 16); Diludia Vates, Butl. from India, &c. (Pl. XCI., Fig. 18); Nephele Hespera, Fabr., from India, Australia, &c. (Pl. XCI., Fig. 20); Acherontia Morta, Hübn., from Java, China, India, &c. (Pl. XCII., Fig. 9); and A. Medusa, Butl., from nearly the same localities as the last (Pl. XCII., Fig. 10). Most of these species fall under Dr. Weismann’s general remarks, so that it is unnecessary to give detailed descriptions. The most divergent marking is that of P. Cyrtolophia, which has a broad white dorsal line bordered with pink, and two large pink ovals on the back of the four anterior segments, the hindmost and larger of these being bisected by the dorsal line. In N. Hespera the subdorsal line is present on segments 6 to 11 only, and it is highly significant that the oblique stripes are absent from these segments, but are present on the anterior segments, where the subdorsal line fails. With reference to the larva of A. Atropos, Mr. Mansel Weale states (Proc. Ent. Soc. 1878, p. v.) that in S. Africa the ordinary form feeds generally on Solanaceæ, whilst the darker and rarer variety is found only on species of Lantana. The following species of these subfamilies are figured by Burmeister: Amphonyx Jatrophæ (loc. cit. Pl. XI., Fig. 1); Protoparce (Diludia) Florestan, Cram. (Fig. 2); Sphinx Justiciæ, Walk. (Fig. 3); Protoparce (Diludia) Lichenea, Walk. (Fig. 4); Sphinx (Protoparce) Cingulata, Fabr. (Pl. XII., Fig. 1); and Sphinx Cestri (Fig. 5). All these species have the characteristic Sphinx-like markings. Dilophonota Ello, Linn. (Pl. XII., Fig. 2), is greenish-brown with a yellow subdorsal line, and D. Hippothöon (Fig. 4), yellow with a whitish subdorsal. Neither of these has oblique stripes. D. Œnotrus, Cram. (Fig. 3), has neither stripes nor subdorsal, but is uniform brown above, passing into green beneath. Protoparce Albiplaga, Walk. (Pl. XIII., Fig. 2, also Mérian, [Pl. III]., and Abbot and Smith, I., Pl. XXIV.), pale green with large yellow, black-bordered patches surrounding the spiracles. Pseudosphinx Tetrio, Linn. (Pl. XIII., Fig. 3), and P. Scyron (Fig. 4) are black with broad transverse belts, yellow and white respectively, encircling the middle of each segment. These light bands serve very effectively to break up the uniform surface of the large bodies of these insects, but the whole marking is suggestive of distastefulness. R.M.]

[122] [The species referred to is placed by Butler in Hübner’s genus Hyloicus. R.M.]

[123] [= Ellema Coniferarum, of Butler’s revision. R.M.]

[124] [= Dilophonota Ello of Butler’s revision. R.M.]

[125] “Synopsis of the North American Sphingides.” Philadelphia, 1859.

[126] [The larvæ of many moths which feed on deciduous trees during the autumn and hibernate, are stated to feed on low-growing plants in the spring, before the buds of their food-trees open. On the other hand, low-plant feeders, such as Triphæna Fimbria, &c., are stated to sometimes feed at night in early spring on the buds of trees. The habits and ontogeny of these species are of special interest in connection with the present researches, and are well worthy of investigation. R.M.]

[127] “Neuer Beitrag zum geologischen Beweise der Darwin’schen Theorie.” 1873, Nos. 1 and 2. [This principle, in common with many others which have only been completely worked out of late years, is foreshadowed by Darwin. Thus, he states when speaking of inheritance at corresponding periods of life: “I could give a good many cases of variations (taking the word in the largest sense) which have supervened at an earlier age in the child than in the parent” (“Origin of Species,” 1st ed., 1860, p. 444). In the case of inherited diseases also: “It is impossible to ... doubt that there is a strong tendency to inheritance in disease at corresponding periods of life. When the rule fails, the disease is apt to come on earlier in the child than in the parent; the exceptions in the other direction being very much rarer.” (“Variation of Animals and Plants under Domestication,” 1st ed., 1868, vol. ii., p. 83.) R.M.]

[128] [If the reddish-brown spots on the larva of S. Populi have the protective function assigned to them by Mr. Peter Cameron (Trans. Ent. Soc. 1880, p. 69), it can be readily understood that they would be of service to the insect in the fourth stage, and the backward transference of this character might thus be accelerated by natural selection, in accordance with the above principles. (See, also, note [100], p. [241].) R.M.]

[129] [For cases of correlation of habit with protective resemblance in larvæ, see a paper in “Ann. and Mag. of Nat. Hist.,” Feb., 1878, pp. 159, 160. Also Fritz Müller on a Brazilian Cochliopod larva, Trans. Ent. Soc. 1878, p. 223. Mr. Mansel Weale states, with reference to S. African Sphingidæ (Proc. Ent. Soc. 1878, p. vi.), that many species when seized “have a habit of doubling up the body, and then jumping a considerable distance with a spring-like action. This is especially the case with species having eye-like markings; and it is probable that if attacked by birds in a hesitating manner, such species might effect their escape amid the grass or foliage.” Many of the defensive weapons and habits of larvæ are doubtless means of protection from ichneumons and other parasitic foes. In the case of saw-flies, Mr. Peter Cameron has shown (Trans. Ent. Soc. 1878, p. 196) that the lashing about of the posterior part of the body may actually frighten away such enemies. The grotesque attitude and spider-like appearance and movements of the caterpillar of Stauropus Fagi are considered by Hermann Müller (“Kosmos,” Nov., 1879, p. 123) to be means of protection from ichneumons. Among the most remarkable means of defence possessed by larvæ is that of secreting a liquid, which Mr. W. H. Edwards has shown, in the case of certain North American Lycænidæ (“Canadian Entomologist.” vol. x., 1878, pp. 3–9 and 131–136), to be attractive to ants, who regularly attend these caterpillars, in the same manner and for the same purpose as they do our aphides. The mutual advantage derived by the ants and larvæ was discovered in the case of Lycæna Pseudargiolus. Mr. Edwards states that the mature larva of this species is singularly free from Hymenopterous and Dipterous parasites:—“Why this species, and doubtless many other Lycænæ, are thus favoured will, perhaps, in some degree appear from a little incident to be related. On 20th June, in the woods, I saw a mature larva on its food-plant; and on its back, facing towards the tail of the larva, stood motionless one of the larger ants.... At less than two inches behind the larva, on the stem, was a large ichneumon-fly, watching its chance to thrust its ovipositor into the larva. I bent down the stem, and held it horizontally before me, without alarming either of the parties. The fly crawled a little nearer and rested, and again nearer, the ant making no sign. At length, after several advances, the fly turned its abdomen under and forward, thrust out its ovipositor, and strained itself to the utmost to reach its prey. The sting was just about to touch the extreme end of the larva, when the ant made a dash at the fly, which flew away, and so long as I watched—at least five minutes—did not return. The larva had been quiet all this time, its tubes out of sight, and head buried in a flower-bud, but the moment the ant rushed and the fly fled, it seemed to become aware of the danger, and thrashed about the end of its body repeatedly in great alarm. But the tubes were not protruded, as I was clearly able to see with my lens. The ant saved the larva, and it is probable that ichneumons would in no case get an opportunity to sting so long as such vigilant guards were about. It strikes me that the larvæ know their protectors, and are able and willing to reward them. The advantage is mutual, and the association is friendly always.” Those who are familiar with Mr. Belt’s description of the standing armies of ants kept by the “bull’s-horn thorn” (“Naturalist in Nicaragua,” pp. 218–222) and by certain Cecropiæ and Melastomæ, will be struck with the analogy between these and the foregoing case. R.M.]

[130] [The adaptive resemblance is considerably enhanced in Catocala and in Lasiocampa Quercifolia by the row of fleshy protuberances along the sides of these caterpillars, which enables them to rest on the tree trunks by day without casting a sharp shadow. The hairs along the sides of the caterpillar of Pæcilocampa Populi doubtless serve the same purpose. (See a paper by Sir John Lubbock, Trans. Ent. Soc. 1878, p. 242; also Peter Cameron, ibid., 1880, p. 75.) It is well known to collectors that one of the best methods of finding the caterpillars of the Catocalæ is to feel for them by day on the barks of their respective food-trees, or to beat for them at night. R.M.]

[131] [See Wallace’s “Contributions to the Theory of Natural Selection,” 1st ed., p. 62. Also a paper in “Ann. Mag. Nat. Hist.” Feb. 1878, p. 159, for cases in point. Rösel in 1746 mentioned this habit in Calocampa Exoleta. Hermann Müller has recorded many other similar instances on the authority of Dr. Speyer; see “Kosmos,” Nov., 1879, p. 114. R.M.]

[132] [Andrew Murray called attention to this fact in 1859 (“Edinburgh New Philos. Journ.,” Jan., 1860, p. 9). This view is also corroborated by the fact that no internal feeders are green; see note [142], p. [310] and Proc. Zoo. Soc. 1873, p. 159. R.M.]

[133] [Proc. Ent. Soc. March 4th, 1867; and “Contributions to the Theory of Natural Selection,” 1st ed., pp. 117–122; also Darwin’s “Descent of Man,” 2nd ed., p. 325. Among the most important recent additions to the subject of the colours, spines, and odours of caterpillars, I may call attention to a paper by Fritz Müller (“Kosmos,” Dec., 1877), the following abstract of which I communicated to the Entomological Society (Proc. 1878, pp. vi, vii):—“The larvæ of Dione Juno and Acræa Thalia live gregariously, and are brown in colour; they are covered with spines, but, being of dull colours, their spiny protection (which in the case of D. Juno is very imperfect) would not preserve them unless they were distinguished as inedible at the right time, and not after being seized, in accordance with the principles laid down by Mr. Wallace. It is suggested that the social habits of the larvæ which lead then to congregate in large numbers, make up for their want of colour, since their offensive odour then gives timely warning to an approaching enemy. The caterpillars of Colænis Julia and Dione Vanillæ are equally wanting in bright colours, but are solitary in their habits, and these species rest on the under side of the leaf when feeding. On the other hand, the caterpillars of Heliconius Eucrate, Colænis Dido, and C. Isabella, which are of solitary habits, and which freely expose themselves, are very gaudily coloured, and therefore most conspicuous. As examples of nearly allied larvæ, of which some species are gregarious and others solitary, Fritz Müller mentions Morpho and Brassolis, which are gregarious; while Opsiphanes and Caligo are solitary. The larva of Papilio Pompeius also is gregarious, and those of P. Nephalion, P. Polydamas, and P. Thoas are solitary.... Fritz Müller sums up his observations by remarking that those caterpillars which live alone, and lack the bright colouring as a sign of offensiveness, must hide themselves; as those of C. Julia and D. Vanillæ. The spiny covering is much less a protection against birds than against smaller enemies; and they may, by the protective habit of living together, diffuse around themselves an offensive atmosphere, even to man, and thus gradually becoming shorter (as with D. Juno), the spines of these caterpillars become useless, and finally are altogether dropped.” See also Sir John Lubbock’s “Note on the Colours of British Caterpillars,” Trans. Ent. Soc. 1878, p. 239. Mr. Peter Cameron finds (Trans. Ent. Soc. 1880, pp. 71 and 75) that these remarks are also applicable to the larvæ of certain saw-flies. In 1877 Mr. J. W. Slater published a paper “On the Food of gaily-coloured Caterpillars” (Trans. Ent. Soc. 1877, p. 205), in which he suggested that such caterpillars might derive their distasteful qualities from feeding on plants containing poisonous or otherwise noxious principles. A much larger number of observations will be required, however, before this view can be accepted as of general application. A beautiful illustration of the theory of warning colours is given by Belt in his “Naturalist in Nicaragua,” p. 321. All the frogs found in the woods round St. Domingo are, with one exception, protectively coloured; they are of nocturnal habits, and are devoured by snakes and birds. The exception was a species of bright red and blue colours, which hopped about by day and made no attempt at concealment. From these facts Mr. Belt concluded that this species was inedible, and on trying the experiment with ducks and fowls this was found to be the case. R.M.]

[134] See the essay “Über den Einfluss der Isolirung auf die Artbilding.” Leipzig, 1872, p. 22.

[135] [See also preceding note [133], p. [294]. R.M.]

[136] [Eng. ed. The habit of hiding by day occurs also in those caterpillars which resemble the bark of their food-trees. Thus Catocala Sponsa and Promissa conceal themselves by day in crevices of the bark, and are, under these circumstances, only found with difficulty. Dr. Fritz Müller also writes to me that in Brazil the caterpillars of Papilio Evander rest in this manner in large numbers, crowded together into dense masses, on the trunks of the orange-trees, which they resemble in colour.]

[137] “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872, p. 21.

[138] I am unfortunately not able to give exact numbers showing the relative proportions of the different forms, since I have never bred S. Convolvuli from eggs, nor C. Elpenor in sufficient numbers.

[139] [With reference to C. Porcellus, see note [71], p. [188]. R.M.]

[140] [In the class of cases treated of in the foregoing portions of this essay, the external conditions remain unaltered during the lifetime of the caterpillar, but change of habit, and in some cases of colour, occurs when the insect has attained a size conceivable à priori, and are realized by observation, in which the environment itself may undergo change during the lifetime of the individual caterpillar. Thus, in the case of hibernating species, the colour which is adaptive to the autumnal colours of the foliage of their food-trees would not assimilate to that of the newly-opened leaves in the spring. I have already quoted (Proc. Zoo. Soc. 1873, p. 155) as instances of what may be called “seasonal adaptation,” the larvæ of Geometra Papilionaria, Acidalia Degenararia, and Gnophos Obscurata, and many more could be named. These species undergo a change of colour before or after hibernation, the change being always adaptive to the environment.

It has long been known that caterpillars which feed on flowers or on plants of variously-coloured foliage, in some cases partake of the colour of their food. See, for instance, Dr. L. Möller’s memoir, “Die Abhängigkeit der Inseckten von ihrer Umgebung,” 1867, and B. D. Walsh “On Phytophagic Varieties and Phytophagic Species,” Proc. Ent. Soc. Philadelph., vol. iii., p. 403. In 1865 Mr. R. McLachlan published a paper entitled “Observations on some remarkable varieties of Sterrha Sacraria, Linn., with general notes on variation in Lepidoptera” (Trans. Ent. Soc. 1865, p. 453), in which he gave many illustrations of this phenomenon. The larva of Heliothis Peltiger, according to Mr. Reading’s description (Newman’s “British Moths,” p. 438), is another case in point. In 1874 a number of instances were published by Mr. Thomas G. Gentry in a paper entitled “Remarkable Variations in Coloration, Ornamentation, &c., of certain Crepuscular and Nocturnal Lepidopterous Larvæ” (“Canadian Entomologist,” vol. vi., p. 85. See also W. H. Edwards’ description of the summer and autumnal larvæ of Lycæna Pseudargiolus; Ibid., vol. x., pp. 12, 13).

The caterpillars of the Sphingidæ appear also in some cases to vary in a manner very suggestive of phytophagic influences. The observations upon S. Ocellatus recorded in the previous [note] (p. [241]) may perhaps be interpreted in this sense. In order to get experimental evidence upon this subject, I may add that Mr. E. Boscher was good enough at my request to repeat his observations, and conduct some breeding experiments during the present year (1880). In the same locality as that previously mentioned, seven larvæ were found feeding on Salix viminalis, all of which were the bright green spotted variety; and in the same osier-bed six more were found on another species of Salix, two of these being the bluish-green variety, and the other four the bright green form. Unless we have here a local race, these observations, in connection with those of last year, tend to show that the light green form is associated with Salix viminalis. When found in the natural state feeding on apple, the caterpillar of this species is generally, perhaps invariably, the bluish-green form. In order to try the effect of breeding the larvæ ab ovo on distinct food-plants, a large number of eggs laid by a female Ocellatus in July were divided into three batches, one being supplied with Salix triandra, another with S. viminalis, and the third lot with apple. The experiment unfortunately failed in great part, owing to most of the larvæ dying off, three from the third lot only surviving; but these were all of the bluish-green form, which colour was shown by all the caterpillars of this batch from their earliest stage. The observation is thus so far successful, as it goes to support the view that the variety mentioned is associated with apple (and S. triandra?) My friend Mr. W. J. Argent informs me that he had a number of specimens of Sphinx Ligustri in his possession this autumn, some of which had been found on lilac and others on laurestinus, and he states that all those on the latter plant had the ground-colour distinctly darker than in those feeding on lilac. I learn also from Mr. W. Davis, of Dartford, that he found a number of these larvæ this year feeding on ash, and that they were all differently coloured to those found on lilac or privet, being of a more greyish-green. Another case of colour-variation in larvæ is that Emmelesia Unifasciata, specimens of which I have recently had an opportunity of examining, through the courtesy of Mr. W. Davis. This species feeds on the seeds of a species of Bartsia when the capsules are in various stages of growth, and (omitting details of marking) those caterpillars found on the green capsules were green, whilst those on the brown capsules were of a corresponding colour.

On the whole I am inclined to believe that sufficient importance has not hitherto been given to phytophagic variability as a factor in determining larval coloration, and a large field for experimental investigation here lies open for future work. The obscure chemico-physiological processes which may perhaps be shown by such researches to lead to phytophagic variation, cannot, I am persuaded, produce any great divergence of character if unaided; but when such causes of variability play into the hands of natural selection variations of direct protective advantage to the species, we can easily see that this all-important agency would seize upon and perpetuate such a power of adaptability to a variable environment. (See Proc. Zoo. Soc. 1873, p. 158, and “Nature,” vol. xiv., pp. 329 and 330.) R.M.]

[141] [In 1879 Mr. George Francis, of Adelaide, forwarded from the latter place a number of moths (a species of Anapæa) together with their larvæ (in alcohol) and cocoons (Proc. Ent. Soc. 1879, p. xvi), and in an accompanying note he stated that the male larva when living is of “a bright emerald green, with red and pink markings on the back, and yellow, black, and white streaks on the sides.” The male larva is described as being smaller than the female, and as possessing all the brilliant colours, the latter “having no red markings, but only white, yellow, and green, with a little black.” I was at first disposed to think that we might be dealing here with two distinct species having differently marked larvæ; but Mr. Francis this present year (1880) forwarded a large number of the living cocoons of this species, which I separated according to size, and, on the emergence of the moths (August), I found that all those from the small cocoons were males, and those from the larger cocoons females. There can be no doubt, therefore, that we have but one species in this case, the larva of which presents the remarkable phenomenon of sexual difference of coloration. As an analogous fact I may here mention the well-known case of Orgyia Antiqua, the larva of which differs in the colour of the tufts of hair according to sex. R.M.]

[142] [I have already given reasons for suspecting that the colour of green caterpillars may be due to the presence of chlorophyll (or some derivative thereof) in their tissues (see Proc. Zoo. Soc. 1873, p. 159). This substance appears to be one of great chemical stability, and, according to Chautard, who has detected it in an unaltered state in the tissues of certain leaf-feeding insects by means of its absorption spectrum (“Comp. Rend.” Jan. 13th, 1873), it resists the animal digestive processes (Ann. Ch. Phys. [5], iii., 1–56). If this view should be established by future observations, we must regard the green colour of caterpillars as having been produced, when protective, from phytophagic variability by the action of natural selection; and the absence of colour in internal feeders, above referred to, is only secondarily due to the exclusion of light, and depends primarily on the absence of chlorophyll in their food. In connection with this I may adduce the fact, that some few species of Nepticula (N. Oxyacanthella, N. Viscerella, &c.) are green, although they live in leaf-galleries where this colour can hardly be of use as a protection; but their food (hawthorn and elm) contains chlorophyll. See also note [130], p. [293]. Further investigations in this direction are much needed. R.M.]

[143] [The same applies to Pseudoterpna Cytisaria, also feeding on broom at the same time of the year. The most striking cases of adaptive resemblance brought about by longitudinal stripes are to be found among fir and pine feeders, species belonging to the most diverse families (Hyloicus Pinastri, Trachea Piniperda, Fidonia Piniaria, &c., &c.) all being most admirably concealed among the needle-shaped leaves. R.M.]

[144] The geographical distribution of the dark form indicates that in the case of this species also, the form referred to is replacing the yellow (green) variety. Whilst in the middle of Europe (Germany, France, Hungary) the dark form is extremely rare, in the south of Spain this variety, as I learn from Dr. Noll, is almost as common as the yellow one. I hear also from Dr. Staudinger that in South Africa (Port Natal) the dark form is somewhat the commoner, although the golden-yellow and, more rarely, the green varieties, occur there. I have seen a caterpillar and several moths from Port Natal, and these all agree exactly with ours. The displacement of the green (yellow) form by the dark soil-adapted variety, appears therefore to proceed more rapidly in a warm than in a temperate climate. [Eng. ed. Dr. Noll writes to me from Frankfort that the caterpillar of Acherontia Atropos in the south of Spain does not, as with us, conceal itself by day in the earth, but on the stems underneath the leaves. “At Cadiz, on the hot, sandy shore, Solanum violaceum grows to a height of three feet, and on a single plant I often found more than a dozen Atropos larvæ resting with the head retracted. It can easily be understood why the lateral stripes are blue when one has seen the south European Solaneæ, on which this larva is at home. Solanum violaceum is scarcely green: violet tints alternate with brown, green, and yellow over the whole plant, and between these appear the yellow-anthered flowers, and golden-yellow berries of the size of a greengage. Thus it happens that the numerous thorns, an inch long, between which the caterpillar rests on the stem, pass from violet into shades of blue, red, green, and yellow.”]

[145] [For Mr. J. P. Mansel Weale’s remarks on the habits of certain ocellated S. African Sphinx-larvæ see note [129], p. [290]. R.M.]

[146] [Some experiments with the caterpillar of C. Elpenor, confirming these results, have been made by Lady Verney. See “Good Words,” Dec. 1877, p. 838. R.M.]

[147] [The eye-spots on Ch. Nerii have thus been supposed by some observers to be imitations of the flowers of the periwinkle, one of its food-plants. See, for instance, Sir John Lubbock’s “Scientific Lectures,” p. 51. R.M.]

[148] “On Insects and Insectivorous Birds,” Trans. Ent. Soc. 1869, p. 21.

[149] Ibid., p. 27.

[150] [Messrs. Weir and Butler inform me that they have not experimented with Sphinx-larvæ. R.M.]

[151] [It appears that the nauseous character of these last butterflies is to a certain extent retained after death, as I found that in an old collection which had been destroyed by mites, the least mutilated specimens were species of Danais and Euplæa, genera which are known to be distasteful when living, and to serve as models for mimicry. See Proc. Ent. Soc. 1877, p. xii. R.M.]

[152] [This bears out the view expressed in a previous note [129], p. [290], that the grotesque attitude and caudal tentacles are more for protection against ichneumons than against larger foes. R.M.]

[153] These experiments, as already mentioned above, were not made with the common German lizard (Lacerta Stirpium), but with the large South European Lacerta Viridis.

[154] Thus, Boisduval states of this caterpillar, which in Provence lives on Euphorbia esula and allied species:—“Its resemblance to a serpent, and its brilliant colour, permit of its being easily discovered.” This was written in 1843, long before natural selection was thought of.

[155] Or some other extinct analogously-marked species.

[156] [See Darwin’s remarks on the struggle for life being most severe between individuals and varieties of the same species “Origin of Species,” 6th ed. p. 59. R.M.]

[157] [Compare this with Darwin’s remarks on “analogous variations,” “Origin of Species,” 6th ed., p. 125. R.M.]

[158] “Zoologische Studien auf Capri. II. Lacerta muralis cærula, ein Beitrag zur Darwin’schen Lehre.” Leipzig, 1874. [The subject of colour-variation in lizards has been much discussed in “Nature” since the publication of the above mentioned essay; see vol. xix., pp. 4, 53, 97, and 122, and vol. xx., pp. 290 and 480. R M.]

[159] “Über die Berechtigung der Darwin’schen Theorie.” Leipzig, 1868. See also the previous essay “On the Seasonal Dimorphism of Butterflies,” pp. [112–116].

[160] [Mr. A. G. Butler has recently advanced the view that this family is not allied to the Sphingidæ, but is related on the one side to the Pyrales, and on the other to the Gelechiidæ. See his paper “On the Natural Affinities of the Lepidopterous Family Ægeriidæ,” Trans. Ent. Soc. 1878, p. 121. R.M.]

[161] I am indebted to my esteemed colleague, Prof. Gestäcker, for the knowledge of this specimen.

[162] Cat. Lep. East India Co., [Pl. VIII].

[163] Such a residue is distinctly visible in S. Ocellatus: see Fig. 70, [Pl. VII].

[164] [The question here also suggests itself as to why the dorsal line should not have been the primary longitudinal stripe, seeing that such a marking is almost naturally produced in many caterpillars by the food in the alimentary canal; or, in other words, why has not natural selection taken advantage of such an obvious means of producing a stripe in cases where it would have been advantageous? In answer to this I may state, that in large numbers of species the dorsal line has thus become utilized; but in the case of large caterpillars resting among foliage, it can be easily seen that light lateral (i.e. subdorsal) stripes, are more effective in breaking the homogeneity of the body than a dorsal line only slightly darker than the general ground-colour. Lateral lines are in fact visible from two directions of space. If a caterpillar thus marked be placed on a twig, these lines are visible when we look at the creature’s back or at either side. That the subdorsal are therefore the primary lines, as shown by Dr. Weismann’s observations of the ontogeny of many of the Sphingidæ, is quite in harmony with the view of their having been produced by natural selection. R.M.]

[165] “Die Darwin’sche Theorie. Elf Vorlesungen über die Entstehung der Thiere und Pflanzen durch Naturzüchtung.” 2nd ed., Leipzig, 1875, p. 195.

[166] [In the following species, already mentioned in previous notes, the oblique stripes are bounded at their upper extremities by a conspicuous subdorsal line:—Acosmeryx Anceus, Cram.; Sphinx Cingulata, Fabr.; Pachylia Ficus, Linn.; P. Syces, Hübn. In Pseudosphinx Cyrtolophia, Butl., the oblique white stripes, beautifully shaded with pink, run into the white pink-bordered dorsal line, so that when seen from above the markings present the appearance of the midrib and lateral veins of a leaf, and are probably specially adapted for this purpose. R.M.]

[167] [The dorsal line as well as the oblique stripes is present in the caterpillar of Smerinthus Tartarinovii, Ménét.; and in Ambulyx Gannascus, Stoll., the oblique stripes are bounded above by a subdorsal line, as in the species named in the preceding note. R.M.]

[168] Cat. Lep. East India Co., Pl. XI.

[169] [Compare this with Darwin’s “Origin of Species” (1st. ed. p. 440), where it is stated that when an animal, during any part of its embryonic career, is active, and has to provide for itself, “the period of activity may come on earlier or later in life; but whenever it comes on, the adaptation of the larva to its conditions of life is just as perfect and beautiful as in the adult animal. From such special adaptations the similarity of the larvæ or active embryos of allied animals is sometimes much obscured.” R.M.]

[170] [For Fritz Müller’s application of this principle to the case of certain groups of Brazilian butterflies see [Appendix II]. to this Part. R.M.]

[171] [The slight variability in the colour of this pupa, opens up the interesting question of the photographic sensitiveness of this and other species, which is stated to cause them to assimilate in colour to the surface on which the larva undergoes its final ecdysis. Some experiments upon this subject have been recorded by Mr. T. W. Wood, Proc. Ent. Soc. 1867, p. xcix, but the field is still almost unexplored. R.M.]

[172] “Über den Einfluss der Isolirung auf die Artbildung.” Leipzig, 1872, p. 20.

[173] In some instances Deilephila Lineata has also been seen by day hovering over flowers.

[174] It is true that I only reared one brood, but from this fifty specimens were obtained. It would be interesting to know whether this variety of the caterpillar is distributed over the whole of Southern Europe.

[175] In this sense Lubbock says:—“It is evident that creatures which, like the majority of insects, live during the successive periods of their existence in very different circumstances, may undergo considerable changes in their larval organization in consequence of forces acting on them while in that condition; not, indeed, without affecting, but certainly without affecting to any corresponding extent, their ultimate form.”—“Origin and Metamorphoses of Insects,” London, 1874, p. 39.

[176] “Grundzüge der Zoologie,” 1875.

[177] [Lepidopterists are of course aware that even these distinctions are not absolute, as no single character can be named which does not also appear in certain moths. The definition in this case, as in that of most other groups of animals and plants, is only a general one. See, for instance, Westwood’s “Introduction to the Classification of Insects,” vol. ii. pp. 330–332. Also some remarks by C. V. Riley in his “Eighth Annual Report” on the insects of Missouri, 1876, p. 170. With reference to the antennæ as a distinguishing character, see Mr. A. G. Butler’s article in “Science for All,” 1880, part xxvii. p. 65. R.M.]

[178] The genus of Morphinæ, Discophora, possesses hairs very similar to those of the genus Cnethocampa belonging to the Bombycidæ.

[179] [The larvæ of genera 14, Phyciodes, and 35, Crenis, are likewise spiny. See Edwards’ “Butt. of N. Amer.” vol. ii. for figures of the caterpillar of Phyc. Tharos: for notes on the larvæ of Crenis Natalensis and C. Boisduvali see a paper by W. D. Gooch, “Entomologist,” vol. xiv. p. 36. The larvæ of genus 55, Ageronia, are also spiny. (See Burmeister’s figure of A. Arethusa, “Lép. Rép. Arg.” Pl. V. Fig. 4). The larvæ of genus 98, Aganisthos, also appear to be somewhat spiny (see Burmeister’s figure of A. Orion, loc. cit. Pl. V. Fig. 6), and this raises the question as to whether the genus is correctly located in its present position. The larvæ of the following genera figured in Moore’s “Lepidoptera of Ceylon,” parts i. and ii., are all spiny:—6, Cirrochroa (Pl. XXXII.); 7, Cynthia (Pl. XXVI.); 27, Kallima (Pl. XIX.); and 74, Parthenos (Pl. XXIV.). Many species of caterpillars which are spiny when adult appear to be spineless, or only slightly hairy when young. See Edwards’ figures of Melitæa Phaeton, Argynnis Diana, and Phyc. Tharos (loc. cit.) and his description of the larva of Arg. Cybele, “Canad. Entom.” vol. xii. p. 141. The spiny covering thus appears to be a character acquired at a comparatively recent period in the phyletic development. R.M.]

[180] [The larvæ of the 110th genus, Paphia, Fabr. (Anæa, Hübn.) are also smoothed-skinned. See Edwards’ figure (loc. cit. vol. i. Pl. XLVI.) of P. Glycerium. Also C. V. Riley’s “Second Annual Report” on the insects of Missouri, 1870, p. 125. Burmeister figures the larva of a species of Prepona (genus 99) which is smooth (P. Demophon, loc. cit. Pl. V. Fig. 1). The horns on the head of Apatura, &c., may possibly be a survival from a former spiny condition. R.M.]

[181] “Synopsis of the described Lepidoptera of North America.” Washington, 1862.

[182] “Catalog der Lepidopteren des Europäischen Faunengebietes.” Dresden, 1871.

[183] This group of moths (“Schwärmer”) is regarded as of very different extents by systematists; when I here comprise under it only the Sphingidæ proper and the Sesiidæ, I by no means ignore the grounds which favour a greater extension of the group; the latter is not rigidly limited. [The affinities of the Sesiidæ (Ægeriidæ) are by no means clearly made out: it appears probable that they are not related to the Sphingidæ. See note [160], p. [370]. R.M.]

[184] [For Mr. A. G. Butler’s observations on the genus Acronycta, see “Trans. Ent. Soc.” 1879, p. 313; and note [68], p. [169], of the present volume. R.M.]

[185] [The following characters are given in Stainton’s “Manual of British Butterflies and Moths,” vol. i. p. 114:—“Larva of very variable form: at one extreme we find the singular Cerura larvæ, with only fourteen legs, and two long projecting tails from the last segment; at the other extreme we have larvæ with sixteen legs and no peculiarity of form, such as Chaonia and Bucephala; most have, however, the peculiarity of holding the hind segment of the body erect when in repose; generally quite naked, though downy in Bucephala and rather hairy in Curtulu; very frequently there are projections on the back of the twelfth segment.” R.M.]

[186] Encyl. Meth. ix. p. 310.

[187] [The genus Vanessa (in the wide sense) appears to be in a remarkable condition of what may be called phyletic preservation. Thus, the group of species allied to V. C.-album passes by almost insensible steps into the group of butterflies typified by our “Tortoiseshells.” The following is a list of some of the intermediate species in their transitional order:—I.-album, V.-album, Faunus, Comma, California, Dryas, Polychloros, Xanthomelas, Cashmirensis, Urticæ, Milberti, &c. Similarly, our Atalanta and Cardui are connected by a number of intermediate forms, showing a complete transition from the one to the other. The following is the order of the species so far as I am acquainted with them:—Atalanta, Dejeanii, Callirhoë, Tammeamea, Myrinna, Huntera, Terpsichore, Carye, Kershawii, and Cardui. R.M.]

[188] “Prodromus Systematis Lepidopterorum.” Regensburg, 1864.

[189] [The larva of Acherontia Morta, figured by Butler (see note [121], p. 262), possesses the characteristically recurved horn; that of Ach. Medusa figured by the same author, does not appear to possess this character in any marked degree. R.M.]

[190] [See note [97], p. [233]. R.M.]

[191] Loc. cit. Pl. XXV. [This species is referred by Butler to the genus Paonias, Hübn. R.M.]

[192] Abbot and Smith, Pl. XXIX. [Placed by Butler in the genus Cressonia, Grote and Robinson. Abbot and Smith state that this larva is sometimes green. According to Mr. Herman Strecker (Lepidop. Rhopal. and Hetero, Reading, Pa. 1874, p. 54) it feeds upon black walnut (Juglans Nigra), hickory (Carya Alba), and ironwood (Ostrya Virginica). Of the North American species of Smerinthus, the following, in addition to Excæcatus, closely resemble our Ocellatus:—S. (Calasymbolus) Geminatus, Say; (C.) Cerisii, Kirby; and Ophthalmicus, Boisd. In addition to S. (Cressonia) Juglandis, S. (Triptogon) Modesta much resembles our Populi. The larva of Geminatus, according to Strecker, is “pale green, lightest above, with yellow lateral granulated stripes; caudal horn violet; stigmata red. It feeds on the willow.” R.M.]

[193] Cat. Brit. Mus.

[194] [This lengthening of the true legs is mimetic according to Hermann Müller, and causes the anterior portion of the caterpillar to resemble a spider. See note [129], p. [290]. R.M.]

[195] [Certain butterflies appear to be crepuscular, if not nocturnal in their habits. Thus in his “Notes on the Lepidoptera of Natal,” Mr. W. D. Gooch states that he never saw Melanitis, Leda, or Gnophodes Parmeno on the wing by day, but generally during the hour after sunset. He adds:—“My sugar always attracted them freely, even up to 10 or 11 p.m.” Many species of Hesperidæ are also stated to be of crepuscular habits by this same observer. See “Entomologist,” vol xvi. pp. 38 and 40. R.M.]

[196] I only make this assumption for the sake of simplicity, and not because I am convinced that the existing Rhopalocera are actually the oldest Lepidopterous group.

[197] Zeitschrift für wissenschaftl. Zoologie, vol. xx. p. 519.

[198] [See for instance Lubbock’s “Origin and Metamorphoses of Insects,” chap. iii.; and F. M. Balfour’s “Comparative Embryology,” vol. i., 1880, pp. 327—356. This last work contains an admirable résumé of our knowledge of the embryonic development of insects up to the date of publication. R.M.]

[199] Are not the 4th, 11th, and 12th segments destitute of the rudiments of legs as in the larvæ of all existing saw-flies? I might almost infer this from Bütschli’s figures (see for instance Pl. XXV., Fig. 17A).

[200] [The grub-formed Hymenopterous larvæ, like the larvæ of all other holometabolous insects, thus represent an acquired degenerative stage in the development, i.e. an adaptation to the conditions of life at that stage. Bearing in mind the above-quoted observations of Bütschli and the caterpillar-like form of the Terebrantiate group of Hymenopterous larvæ, the following remarks of Balfour’s (loc. cit. p. 353), appear highly suggestive:—“While in a general way it is clear that the larval forms of insects cannot be expected to throw much light on the nature of insect ancestors, it does nevertheless appear to me probable that such forms as the caterpillars of the Lepidoptera are not without a meaning in this respect. It is easy to conceive that even a secondary larval form may have been produced by the prolongation of one of the embryonic stages; and the general similarity of a caterpillar to Peripatus, and the retention by it of post-thoracic appendages, are facts which appear to favour this view of the origin of the caterpillar form.” See also Sir John Lubbock, loc. cit., pp. 93 and 95. R.M.]

[201] [In the most recent works dealing with this order six groups, based on the character of the imagines are recognized, viz.:—Tubulifera, Terebrantia, Pupivora, Heterogyna Fossores, and Mellifera. (See, for instance, F. P. Pascoe’s “Zoological Classification,” 2nd ed. p. 147.) Of these groups the larvæ of the Terebrantia as thus restricted are all of the caterpillar type (Tenthredinidæ and Siricidæ), whilst those of the other groups are maggot-shaped. For a description of the development of the remarkable aberrant larva of Platygaster, see Ganin in Zeit. f. wissenschaftl. Zool., vol. xix. 1869. R.M.]

[202] [For recent investigations on the structure of the thorax in Diptera, see a paper by Mr. A. Hammond, in Journ. Linn. Soc., Zoology, vol xv. p. 9. R.M.]

[203] I am familiar with the fact that the two sub-orders of true Diptera, the short-horned (Brachycera), and the long-horned (Nemocera), are not sharply limited; and I am likewise well acquainted with the circumstance that there are forms which connect the two larval types. The connecting forms of the imagines do not, however, always coincide with the intermediate larval forms, so that there here arises a second and very striking incongruence of morphological relationship which depends only upon the circumstance that the one stage has diverged in form more widely than the other through a greater divergence in the conditions of life. The difficulty is in these cases aggravated because an apparent is added to the true form-relationship through convergence, so that without going into exact details the form and genealogical relationships of the Diptera cannot be distinguished. It would be of great interest for other reasons to make this investigation, and I hope to be able to find leisure for this purpose at some future period.

[204] “Entwicklung der Dipteren.” Leipzig, 1864.

[205] Lubbock concludes from the presence of thoracic legs in the embryonic larva of bees that these have been derived from a larva of the Campodea type, but he overlooks the fact that the rudiments of the abdominal legs are also present; loc. cit., p. 28.

[206] “Für Darwin,” Leipzig, 1864, p. 8.

[207] Mem. Peabody Acad. of Science, vol. i. No. 3.

[208] Verhandl. Wien. Zoolog. Botan. Gesellsch. 1869, p. 310.

[209] Über Ontogenie und Phylogenie der Insekten. Eine akademische Preisschrift. Jen. Zeitschrift. Bd. x. Neue Folge, iii. Heft 2. 1876. [Some remarks by F. M. Balfour on the origin of certain larval forms have already been quoted in a previous [note] (p. [485]). This author further states:—“The fact that in a majority of instances it is possible to trace an intimate connection between the surroundings of a larva and its organization proves in the clearest way that the characters of the majority of existing larval forms of insects have owed their origin to secondary adaptations. A few instances will illustrate this point:—In the simplest types of metamorphosis, e.g. those of the Orthoptera genuina, the larva has precisely the same habits as the adult. We find that a caterpillar form is assumed by phytophagous larvæ amongst the Lepidoptera, Hymenoptera, and Coleoptera. Where the larva has not to go in search of its nutriment the grub-like apodous form is assumed. The existence of such an apodous larva is especially striking in the Hymenoptera, in that rudiments of thoracic and abdominal appendages are present in the embryo and disappear again in the larva.... It follows from the above that the development of such forms as the Orthoptera genuina is more primitive than that of the holometabolous forms, &c.” Comparative Embryology, vol. 1, p. 352. R.M.]

[210] [The Aphaniptera are now recognized in this country as a sub-order of Diptera. See, for instance, Huxley’s “Anatomy of Invertebrated Animals,” p. 425, and Pascoe’s “Zoological Classification,” 2nd ed. p. 122. R.M.]

[211] [This illustration of course only applies to the old arrangement of the Hymenoptera into Terebrantia and Aculeata. See also note [201], p. [488]. R.M.]

[212] [Eng. ed. This law is perhaps a little too restricted, inasmuch as it is theoretically conceivable that the organism may be able to adapt itself to similar conditions of life in different ways; differences of form could thus depend sometimes upon differences of adaptation and not upon differences in the conditions of life, or, as I have formerly expressed it, it is not necessary to allow always only one best mode of adaptation.]

[213] [It must be understood that the word rendered here and elsewhere throughout this work as “transformation” is not to be taken in the narrow sense of metamorphosis, but as having the much broader meaning of a change of any kind incurred by an organism. Metamorphosis is in fact but one phase of transformation. R.M.]

[214] By the Editor.

[215] Mr. C. V. Riley in his excellent “Annual Reports” already quoted in previous notes, states that the larvæ of Agrotis Inermis, Leucania Unipuncta (Army-worm), and L. Albilinea are all loopers when newly hatched. (See First Report, p. 73; Eighth Report, p. 184; and Ninth Report, p. 53.)

[216] The following species not referred to in the previous part of this work are figured by Semper (Beit. zur Entwicklungsgeschichte einiger ostasiat. Schmet.; Verhandl. d. k.k. zoo. bot. Gesell. in Wien, 1867):—Panacra Scapularis, Walk.; Chærocampa Clotho, Drury; and Diludia (Macrosila) Discistriga, Walk. The following are figured by Boisduval and Guenée. (Spéc. Gén. 1874):—Smerinthus Ophthalmicus, Boisd.; Sphinx Jasminearum, Boisd.; S. (Hyloicus) Plebeia, Fabr.; S. (Hyloicus) Cupressi, Boisd.; S. (Pseudosphinx) Catalpæ, Boisd.; Philampelus Jussiuæ, Hübn. (= Sphinx Vitis, Linn.?); and Ceratomia Amyntor, Hübn. As the works of Abbot and Smith, and Horsfield and Moore have been exhausted by Dr. Weismann, it is quite unnecessary to extend this note by giving a list of the species figured by these authors.

[217] The same inference has already been drawn with respect to Pterogon (Proserpinus) Œnotheræ, see pp. [257, 258].

[218] This would of course be the fourth segment if the head be considered the first, as on the Continent.

[219] “Second Annual Report,” 1870, p. 78.

[220] “Entomologist,” vol. xiv. p. 7.

[221] With reference to the habits of C. Capensis (p. [531]), I have since been informed by Mr. Trimen that this species does not conceal itself by day, so that the dimorphism may be regarded as a character retained from an earlier period and adapted to the present life conditions.

[222] “Kosmos,” Dec. 1877, p. 218. The paper is here introduced chiefly with a view to illustrate an important case of incongruence among Lepidopterous pupæ.

[223] [Maracujá, the local name for the Passiflora. R.M.]

[224] See p. [448].

[225] Verhandl. Schweiz. Naturforsch. Gesellschaft. Einsiedeln, 1868.

[226] [Eng. ed. In 1878 Señor José M. Velasco published a paper entitled “Description, metamorfosis. y costumbres de una especie nueva del genero Siredon.” Memor. Sociedad Mexicana de Historia Natural, December 26th. See Addendum to this essay.]

[227] Dana and Silliman’s Amer. Journ., 3rd series, i. p. 89. Annals Nat. Hist. vii. p. 246.

[228] Proc. Zoo. Soc. 1870, p. 160.

[229] Compt. Rend., vol. lx. p. 765 (1865).

[230] Nouvelles Archives du Muséum d’Histoire Nat. Paris, 1866, vol. ii. p. 268.

[231] Proc. Boston Soc., vol. xii. p. 97; Silliman’s Amer. Journ., vol. xlvi. p. 364; reference given in “Troschel’s Jahresbericht” for 1868, p. 37.

[232] Proc. Boston Soc., vol. xii. p. 97; Silliman’s Amer. Journ., vol. xlvi. p. 364. I have not been able to get a copy of this paper, and quote from a reference in “Troschel’s Jahresbericht.” See preceding note.

[233] Dana and Silliman’s Amer. Journ. See note 3.

[234] Proc. Acad. Philadelph. xix. 1867, pp. 166–209.

[235] Mém. Acad. Petersb. vol. xvi.

[236] [Eng. ed. Seidlitz is an exception, since in his work on Parthenogenesis (Leipzig, 1872, p. 13) he states that “In the Axolotl, Pædogenesis, which is not in this case... monogamous, but sexual, and indeed gynækogenetic, has already become so far constant that it has perhaps entirely superseded the orthogenetic reproduction.”]

[237] Über den Einfluss der Isolirung auf die Artbildung. Leipzig, 1872, p. 33.

[238] Duméril represents the teeth of the vomer as separated from those of the os palatinum by a gap. This is probably accidental, since Gegenbaur (Friedrich u. Gegenbaur, the skull of Axolotl, Würzburg, 1849) figures the rows of teeth as passing over from the one bone to the other without interruption. This was the case with the Axolotls which I have been able to examine on this point; but this small discrepancy is, however, quite immaterial to the question here under consideration.

[239] See O. Hertwig “Über das Zahnsystem der Amphibien und seine Bedeutung für die Genese des Skelets der Mundhöhle.” Archiv. für microsc. Anat., vol. xi. Supplement, 1874.

[240] [Eng. ed. These Amblystomas have since died and have been minutely described by Dr. Wiedersheim. See his memoir, “Zur Anatomie des Amblystoma Weismanni,” in Zeit. für wiss. Zool., vol. xxxii. p. 216.]

[241] See Strauch, loc. cit. p. 10.

[242] See [Part I]. of this volume.

[243] [This is the principle of “Degeneration” recognized by Darwin (see “Origin of Species,” 6th ed. p. 389, and “Descent of Man,” vol. i. p. 206), and given fuller expression to by Dr. Anton Dohrn (see his work entitled “Der Ursprung der Wirbelthiere und das Princip des Functionswechsels.” Leipzig, 1875). A large number of cases have been brought together by Prof. E. R. Lankester, in his recent interesting work on “Degeneration, a Chapter in Darwinism.” Nature series, 1880. R.M.]

[244] “Sulla Larva del Triton Alpestris.” Archivio per la Zoologia. Genova e Torino, 1861, vol. i. pp. 206–211.

[245] See also Lubbock “On the Origin and Metamorphoses of Insects,” London, 1874.

[246] See the first essay “On the Seasonal Dimorphism of Butterflies,” p. [82].

[247] [Eng. ed. It has frequently been objected to me that the existing Axolotl is not a form resulting from atavism, but a case of “arrested growth.” The expression “atavism” is certainly to be here taken in a somewhat different sense than, for example, in the case of the reversion of the existing Axolotl to the Amblystoma form. Further on, I have myself insisted that in the first case the phyletic stage in which the reversion occurred is still completely preserved in the ontogeny of each individual, whilst the Amblystoma stage has become lost in the ontogeny of the Axolotl. If, therefore, we apply the term “atavism” only to such characters or stages (i.e. complexes of characters) as are no longer preserved in the ontogeny, we cannot thus designate the present arrest of the Axolotl at the perennibranchiate stage. Such a restriction of the word, however, appears to me but little desirable, since the process is identical in both cases, i.e. it depends upon the same law of heredity, in accordance with which a condition formerly occurring as a phyletic stage suddenly reappears through purely internal processes. It is true that the reversion is not complete, i.e. the present sexually mature Axolotl does not correspond in all details with its perennibranchiate ancestors. Since Wiedersheim has shown that the existing Axolotl possesses an intermaxillary gland, this can be safely asserted. This gland occurs only in land Amphibians, and therefore originated with the Amblystoma form, afterwards becoming transferred secondarily to the larval stage. Nevertheless, the present Axolotl must resemble its perennibranchiate ancestors in most other characters, and we should be the more entitled to speak of a reversion to the perennibranchiate stage as we speak also of the reversion of single characters. To this must be added that the Axolotl does not correspond exactly with an Amblystoma larva, since Wiedersheim has shown that the space for the intermaxillary gland is present, but that the gland itself is confined to a few tubes which do not by any means fill up this space. (“Das Kopfskelet der Urodelen.” Morph. Jahrbuch, vol. iii. p. 149). By the expression “arrested growth” not much is said, if at the same time the cause of the arrest is left unstated. But what can be the cause why the whole organization remains stationary at the perennibranchiate stage, the sexual organs only undergoing further development? Surely only that law or force of heredity known by its effects, but obscure with respect to its causes, through which old phyletic stages sometimes suddenly reappear, or in other words, that power through which reversion takes place. It must not be forgotten that all these cases of “larval reproduction” in Amphibians appear suddenly. The present sexually mature form of the Axolotl has not arisen by the sexual maturity gradually receding in the ontogeny from generation to generation, but by the occurrence of single individuals which were sexually mature in the perennibranchiate stage, these having the advantage over the Amblystomæ in the struggle for existence under changed climatic conditions.

By admitting a reversion, we perfectly well explain why arrest at the perennibranchiate stage can be associated with complete development of the sexual organs; the assumption of an “arrested growth” leaves this combination of characters completely unexplained. Moreover, I am of opinion that the expressions “arrested growth” or “reversion” are of but little importance so long as the matter itself is clear.]

[248] See Haeckel’s “Anthropogenie,” p. 449.

[249] “Der Ursprung der Wirbelthiere und das Princip des Functionswechsels,” Leipzig, 1875.

[250] Bull. Soc. Neuchâtel. vol. viii. p. 192. Reference given in “Troschel’s Jahresbericht” for 1869.

[251] Sitzungsberichte d. math. phys. Klasse der Akad. d. Wiss. zu München, 1875. Heft i.

[252] Compt. Rend. vol. lxviii. pp. 938 and 939.

[253] Archiv f. Naturgeschichte, 1867.

[254] Compt. Rend. vol. v. 1870, p. 70.

[255] Bull. Soc. Neuchâtel. vol. viii. p. 192. Reference given in “Troschel’s Jahresbericht” for 1869.

[256] [Eng. ed. It was mentioned in the German edition of this work that in the spring of 1876 a female Amblystoma of the Jardin des Plantes in Paris had laid eggs (see Blanchard in the Compt. Rend. 1876, No. 13, p. 716). Whether these eggs were fertile, or whether they developed was not then made known. Thus much was however at the time clear, that even if this had been the case, the reproduction of this Amblystoma would have been only an exceptional occurrence. At that time there were in the Jardin des Plantes Amblystomas which had been kept for more than ten years, and only on one occasion was there a deposition of eggs, and this by only one specimen. That I was correct in speaking of the “sterility” of these Amblystomas in spite of this one exception, is proved by the latest communication from the Jardin des Plantes. We learn from this (Compt. Rend. No. 14, July, 1879, p. 108) that in the years 1877 and 1878 none of the Amblystomas laid any more eggs, although all means were exerted to bring about propagation. In April, 1879, eggs were again laid by one female, and by a second in May. These eggs certainly developed, as did those of 1876, and produced tadpoles. These Amblystomas are therefore not absolutely, but indeed relatively sterile. Whilst the Axolotl propagates regularly and freely every year, this occurs with the Amblystoma but rarely and sparsely. The degree of their sterility can only be approximately established when we know the number of Amblystomas that have since been kept in the Jardin des Plantes. Unfortunately nothing has been said with respect to this.]

[257] Origin of Species, 6th ed. p. 252.

[258] In plants also reversion forms show sterility in different degrees. Mr. Darwin has called my attention to the fact that the peloric (symmetrical) flowers which occasionally appear as atavistic forms in Corydalis solida are partly sterile and partly fertile. That in other causes of sterility, and above all by bastardizing, the reproductive power is lost in the most varying degrees, has been known since the celebrated observations of Kölreuter and Gärtner. [Eng. ed. An Orchid (Catasetum tridentatum) has the sexes separate, and the male flowers (Myanthus barbatus) differ considerably from the female (Monachanthus viridis); besides these, there occurs a form with bisexual flowers which must be considered as a reversion (Cat. tridentatum) and this is always sterile. Darwin, “Fertilization of Orchids,” 2nd ed. p. 199.]

[259] As we do not know the origin of the “Paris Axolotl” I must restrict myself in the following remarks to Siredon Mexicanus (Shaw).

[260] Mühlenpfordt, “Versuch einer getreuen Schilderung der Republik Mejico,” Hanover, 1844, vol. ii. p. 252.

[261] [The specific gravity of sea water (Atlantic), according to the determinations of Mr. Buchanan on board the “Challenger,” at 15.56° C. varies from 1.0278 to 1.0240. That of the water of the Dead Sea is 1.17205.—Watts’ “Dict. of Chemistry,” vol. v., table, p. 1017. R.M.]

[262] Loc. cit. p. 252.

[263] “Über die specifische Verschiedenheit des gefleckten und des schwarzen Erdsalamanders oder Molchs, und der höchst merkwürdigen, ganz eigenthümlichen Fortpflanzungsweise des Letzteren.” Isis, Jahrg. 1833, p. 527.

[264] The experiments referred to have not been made known; I am indebted for them to a written communication kindly furnished by an esteemed colleague.

[265] See Mühlenpfordt’s work already quoted, vol. i.

[266] In the province of botany such a case has already been made known by Fritz Müller (Botan. Zeitung, 1869, p. 226; 1870, p. 149). I may be here permitted to quote a passage from the letter in which Dr. Müller calls attention to this interesting discovery. “As a proof of the possibility that a reversion form can again become a persistent character in a species or in the allied form of a particular district, I may refer you to an Epidendrum of the island of Santa Catharina. In all Orchids (with the exception of Cypripedium) only one anther is developed; in very rare cases well-formed anthers appear as reversions among the aborted lateral anthers of the inner whorl. In the Epidendrum mentioned, these are however always present.”

[267] [This species is interesting as being ovoviviparous, the young passing through the branchiate stage within the body of the mother. Some experiments, which were partially successful, were made by Fräulein v. Chauvin with a view to solve the question whether the branchiate stage could be prolonged by taking the larvæ directly from the mother before birth and keeping them in water. See “Zeit. für wissen. Zoo.” vol. xxix., p. 324. R.M.]

[268] See Fatiot, “Les Reptiles et les Batraciens de la haute Engadine.” Geneva, 1873.

[269] I can remember at Upper Engadine a peculiar kind of preserved beef, prepared by simply drying in the air; also the mummification of entire human bodies by drying in the open air, as is practised at Great St. Bernard.

[270] “Faune des Vertébrés de la Suisse,” vol. iii. “Histoire Naturelle des Reptiles et des Batraciens.” Geneva, 1873.

[271] See Wiedersheim, “Versuch einer gleichenden Anatomie der Salamandrinen.” Würzburg, 1875.

[272] See Gené, “Memorie della Reale Acad. di Torino,” vol. i.

[273] Rana esculenta never reaches Alpine regions, this species not having been found higher than 1100 meters. (Fatiot, loc. cit., p. 318.)

[274] See also the excellent work upon Mexico by Mühlenpfordt already quoted, vol. i., pp. 69–76.

[275] “Essai politique sur le Royaume de la Nouvelle Espagne,” 1805, p. 291.

[276] [The expression made use of by the author, viz. “Diluvialzeit,” would perhaps be more in harmony with the views of English geologists if rendered as the “pluvial period,” thereby indicating the period of excessive rainfall which, according to Mr. Alfred Tylor, succeeded to and was a consequence of the thawing of the great glaciers which accumulated during the last glacial epoch. There is abundant evidence to show that during the latter period glacial action extended in North America at least as far south as Nicaragua. See Belt on “The Glacial Period in North America,” Trans. Nova Scotian Inst. of Nat. Sci. 1866, p. 93, and “The Naturalist in Nicaragua,” pp. 259–265. R.M.]

[277] [Eng. ed. A memoir by Samuel Clarke has since been published upon the embryonic development of Amblystoma punctatum, Baird. Baltimore, 1879.]

[278] [Eng. ed. See this author’s work, “Das Kopfskelet der Urodelen.” Leipzig, 1877, p. 149.]

[279] [See preceding note 52. R.M.]

[280] See note [226], p. [566].

[281] [Prof. Semper also remarks (“Animal Life,” note 47, p. 430) with reference to the Axolotl of Lake Como in the Rocky Mountains, which he states always becomes transformed into Amblystoma Mavortium, that this metamorphosis “takes place in the water, and the Amblystomas, so long as they are little, actually live exclusively in the water, as I know by my own experience. A young Amblystoma which I kept alive for a long time, never went out of the water of its own free will, while one nearly twice as large lives entirely on land and only takes a bath now and then. It always goes into the water when the temperature of the air in the cellar, in which my aquaria stand, falls below that of the water—down to about 6° or 8° C.” This statement appears to suggest that the effect of temperature may be a factor in some way concerned in these interesting cases of transformation, and would in any case be well worthy of experimental investigation. Some further details concerning the Siredon Lichenoides of Lake Como have been recently published by Mr. W. E. Carlin (Proc. U.S. National Museum, June, 1881). The lake, which is shallow, is fed by a constant stream of fresh water, but the water of the lake is intensely saline. The Siredon never enter the fresh water stream, but congregate in large numbers in the alkaline waters of the lake. “When about one hundred and fifty were placed in fresh water they seemed to suffer no inconvenience, but it had a remarkable effect in hastening their metamorphosis into the Amblystoma form. Of an equal number kept in fresh water and in the lake water, quite a change occurred with the former after twenty-four hours, while the latter showed no change after several days of captivity. Those that were kept well fed in jars usually began to show a slight change in from two to three weeks, and all of them completed the change into the Amblystoma inside of six weeks, while in some kept, but not specially fed, there were but three changes in three months.” (Nature, Aug. 25th, 1881, p. 388.) R.M.]

[282] [Some experiments on the transformation of the Crustacean Artemia Salina into A. Milhausenii by gradually increasing the saltness of the water, and conversely, the transformation of A. Milhausenii into A. Salina by diminishing the saltness of the water, have been made by Schmankewitsch (Zeitschrift f. wiss. Zool. xxv. Suppl. 103 and xxix. 429), but the changes which occur here are much less considerable than in the case of the Axolotl. R.M.]

[283] “Reden und kleinere Aufsätze, Th. II.: Studien aus dem Gebiete der Naturwissenschaften.” St. Petersburg, 1876, p. 81.

[284] This obviously does not imply that the naturalist should not investigate Nature’s processes, and not only correlate these, but also work them up into a universal conception; this is indeed both desirable and necessary if natural knowledge is to be regarded in its true value. The naturalist by this means becomes a philosopher, and the vitality of the so-called “natural philosopher” has been inspired, not by the necessity for investigation, but by philosophy proper.

[285] [The discovery here referred to is the synthesis of urea by Wöhler in 1828 (Pogg. Ann. xii., 253; xv. 619), by the molecular transformation of ammonium cyanate. Since that period large numbers of organic syntheses have been effected by chemists, and many of the compounds formerly supposed to be essential products of life have been built up in the laboratory from their inorganic elements. The division of chemistry into “organic” and “inorganic” is thus purely artificial, and is merely retained as a matter of convenience, the former division of the science being defined as the chemistry of the carbon compounds. R.M.]

[286] “Wahreit und Irrthum im Darwinismus.” Berlin, 1875.

[287] [Eng. ed. I have been reproached by competent authorities for having clothed my ideas upon the theory of selection in the form of a reply to Von Hartmann. I willingly admit that this author cannot be considered as the leader of existing philosophical views upon the theory of descent in Germany; Frederick Albert Lange has certainly a much greater claim to this position. Lange does not however combat this theory; he accepts and develops it most beautifully and lucidly on a sound philosophical basis in such a manner as has never been done before from this point of view (“Geschichte des Materialismus,” 3rd. ed., 1877, vol. ii. pp. 253–277). On most points I can but agree with Lange. Von Hartmann, however, whose objections appeared to me to be supported by a wide scientific knowledge, afforded me a suitable opportunity of developing my own ideas upon some essential points in the theory of selection. In this sense only have I attempted to interfere with this author, the refutation of his views, as such, having been with me a secondary consideration.] [The chief exponent of the doctrine of organic evolution in this country is Mr. Herbert Spencer, in whose “Principles of Biology,” vol. i. chap. xii., will be found a masterly treatment of the theory of descent from a “mechanical” point of view. R.M.]

[288] [The above views on the nature of variability, which were also broadly expressed in the first essay “On the Seasonal Dimorphism of Butterflies” (pp. [114], 115), are fully confirmed by Herbert Spencer (loc. cit. chaps. ix. and x.), and more recently by A. R. Wallace in an article on “The Origin of Species and Genera” (Nineteenth Century, vol. vii., 1880, p. 93). See also some remarks by Oscar Schmidt in his “Doctrine of Descent and Darwinism,” Internat. Scien. Ser. 3rd. ed. 1876, p. 173. R.M.]

[289] [This law has been beautifully applied by Herbert Spencer in order to explain why, with an unlimited supply of food, an organism does not indefinitely increase in size. “Principles of Biology,” vol. i. p. 121–126. R.M.]

[290] [Eng. ed. This idea, formerly expressed by me, occurs also in Lange (“Geschichte des Materialismus,” ii. 265), and is there exemplified in a very beautiful manner by illustrations from modern chemistry. Lange compares what I have termed above the “physical constitution” of the organism to the chemical constitution of one of those organic acids which by substitution of single elements may become transformed into more complicated acids, but which, as it were, always undergo “further development” in only one determined and narrowly restricted course. Here, as with the organism, the number of possible variations is very great, but is nevertheless limited, since “what can or cannot arise is determined beforehand by certain hypothetical properties of the molecule.”]

[291] “Origin of Species.” 4th German ed., p. 19; 5th English ed., p. 6.

[292] [Mr. A. R. Wallace, in his article last referred to, quotes some most valuable measurements of mammals and birds, showing the amount of variation of the different parts. These observations were published by J. A. Allen, in a memoir “On the Mammals and Winter Birds of East Florida,” &c. (Bulletin of the Museum of Comparative Zoology at Harvard College, Cambridge, Mass., vol. ii. No. 3.) R.M.]

[293] [See note [142], p. [310]. R.M.]

[294] “Die Darwin’sche Theorie,” Dorpat, 1875.

[295] [A certain number of instances of mimicry are known to occur between species both of which are apparently nauseous. A most able discussion of this difficult problem is given by Fritz Müller, in the case of the two butterflies Ituna Ilione and Thyridia Megisto, in a paper published in Kosmos, May, 1879 (p. 100). The author shows by mathematical reasoning that such resemblances between protected species can be accounted for by natural selection if we suppose that young birds and other insect persecutors have to learn by experience which species are distasteful and which can be safely devoured. See also Proc. Ent. Soc. 1879, pp. xx-xxix. R.M.]

[296] See Haeckel’s “Generelle Morphologie,” ii. 107.

[297] “Über die Berechtigung der Darwin’schen Theorie,” Leipzig, 1868.

[298] “Populäre wissenschaftl. Vorträge,” vol. ii., Brunswick, 1871, p. 208.

[299] “Das Unbewusste vom Standpunkte der Physiologie u. Descendenztheorie,” Berlin, 1872, p. 89. The second edition appeared in 1877, in Von Hartmann’s own name.

[300] “Über die Berechtigung,” &c., Leipzig, 1868. In this work will be found briefly laid down the theoretical conception of variability here propounded somewhat more broadly. [In the last edition of the “Origin of Species” Darwin states, with respect to the direct action of the conditions of life as producing variability, that in every case there are two factors, “the nature of the organism and the nature of the conditions.” 6th ed. p. 6. R.M.]

[301] [Although hardly necessary to the evolutionist, it may perhaps be well to remind the general reader, that all experiments upon spontaneous generation, or abiogenesis, have hitherto yielded negative results; no life is produced when the proper precautions are taken for excluding atmospheric germs. But although we have so far failed to reproduce in our laboratories the peculiar combination of conditions necessary to endow colloidal organic matter with the property of “vitality,” the consistent evolutionist is bound to believe, from the analogy of the whole of the processes of nature, that at some period of the earth’s history the necessary physical and chemical conditions obtained, and that some simple form or forms of life arose “spontaneously,” i.e. by the operation of natural causes. R.M.]

[302] See Haeckel’s “Generelle Morphologie,” vol. ii. p. 203, and Seidlitz, “Die Darwin’sche Theorie,” 1875, p. 92 et seq.

[303] [In a recently published work by Dr. Wilhelm Roux this author has attempted to work out the idea of an analogy between the struggle for existence and survival of the fittest in individuals and species, and the struggle for existence and survival of the parts in the individual organism. See “Der Kampf der Theile im Organismus: ein Beitrag zur Vervollständigung der mechanischen Zweckmässigkeitslehre,” Leipzig, 1881. R.M.]

[304] [Eng. ed. Meanwhile it has been shown by Oscar Schmidt that Von Hartmann, under the name of “the Unconscious,” re-invests the old vital force with some portion of its former power. “Die naturwissenschaftlichen Grundlagen der Philosophie des Unbewussten,” Leipzig, 1877, p. 41.]

[305] Loc. cit. p. 175.

[306] Loc. cit. p. 156.

[307] “Über die Cuninen-Knospenähren im Magen von Geryonien.” Reprint from “Mittheil. des naturwiss. Vereines,” Graz, 1875.

[308] [See Darwin’s “Origin of Species,” 6th ed. pp. 33, 34, and 201–204. R.M.]

[309] [Eng. ed. See Kant’s “Allgemeine Naturgeschichte und Theorie des Himmels.”]

[310] “Das Unbewusste vom Standpunke der Physiologie und Descendenz-Theorie,” Berlin, 1872, p. 16.

[311] [Eng. ed. See Lotze’s “Mikrokosmos,” 1st ed., vol. iii. pp. 477–483.]

[312] See Helmholtz’s “Populäre wissenschaftl. Vorträge,” vol. ii., Brunswick, 1872.

[313] See also Fr. Vischer’s “Studien über den Traum. Beilage zur Augsburger Allgem. Zeitung,” April 14th, 1876. Haeckel also includes this idea in his recent essay already quoted, “Die Perigenesis der Plastidule,” Berlin, 1876, p. 38 et seq.

[314] See Von Hartmann, loc. cit. p. 158.

INDEX.

• A.
• Abbot and Smith, descriptions of larvæ, [192], [216], [261], [263], [268], [452], [453].
• Acherontia Atropos, South African form of larva, [263], [324], [531];
• larva in Spain, [324];
• larva phytophagically variable, [531].
• Acosmeryx anceus, larva, [192].
• Acræa and Maracujà butterflies as larvæ, pupæ, and imagines, [536].
• Acronycta, affinities of genus, [169].
• Adaptation, [281], [589];
• analogous, [376];
• physiological, [590];
• mutual, [647].
• Allen, J. A., variation in birds and mammals, [658].
• Alpine form of P. Napi, [40].
• Alpine hare, seasonal change of colour, [7], [660].
• Alternation of generations, [80], [699], [702].
• Amblystoma. A. mavortium from Siredon lichenoides, [567];
• A. tigrinum bred by Duméril, [567];
• distribution of species of genus, [569];
• habits of species, [573];
• a reversion form, [531], [592];
• sterility of, [593];
• oviposition by, [594], [623];
• definition of genus, [610];
• degeneration of, [612];
• causes of reversion of, [600], [608], [613], [620], [631];
• A. punctatum and A. fasciatum, development of, [623];
• summer sleep of, [628];
• A. mavortium, metamorphosis of, [629].
• Ambulyx gannascus and A. liturata, larvæ, [245].
• Amixia, the principle of, [46], [110].
• Ammonites, [275].
• Ampelophaga rubiginosa, larva, [192].
• Amphibia of Upper Engadine, [615].
• Amphiuma, characters of genus, [579].
• Anceryx, larva, [264];
• stages 3, 4, and 5, [266].
• Aphaniptera, characters of, [498].
• Aphides, organic reproduction of, [97].
• Araschnia. A. levana and A. prorsa seasonally dimorphic, [2];
• dimorphism of larvæ, [6];
• A. levana, first experiments with, [10];
• A. var. porima, artificial production of, [10], [17];
• A. levana, Dorfmeister’s experiments with, [11];
• explanation of seasonal dimorphism of, [19];
• northern, extension of, [20];
• A. prorsa gradual origination of, [22];
• A. levana, reversion of, caused by high temperature, [37];
• the levana form the most constant, [43], and sexually dimorphic, [43];
• summer and winter forms compared, [56];
• experiments with A. levana, [117];
• A. levana, variable in all three stages, [405];
• incongruence in genus, [449].
• Argent, W. J., phytophagic variability of larva of Sphinx Ligustri, [306].
• Argynnis paphia, dimorphism of, [250].
• Artemia salina and A. Milhausenii, transformations of, [635].
• Ascaris nigrovenosa, propagation of, [90].
• Askenasy, on limits of variability, [114].
• Atavism and arrested growth, [587].
• Aterica meleagris, colour variations of, [8].
• Atoms, actions of, [710];
• sensibility of, [714].
• Axolotl, transformation into Amblystoma, [555];
• experiments with, [558];
• duration of transformation, [563];
• conditions of metamorphosis of, [564];
• distinguishing characters, [575].
• B.
• Baer, Karl Ernst von, on the mechanism of nature, [639];
• on the vital force, [641];
• on the theory of selection, [694];
• on causality and purpose, [708].
• Baird, Prof., on the development of species of Amblystoma, [623].
• Balbiani, reproduction of Aphides, [97].
• Balfour, F. M., on insect phylogeny, [485], [494].
• Bates, H. W., local variation in Amazonian Lepidoptera, [60].
• Bees, embryological development of, [483].
• Belt, Thomas, on coloured frogs, [294];
• on the glaciation of North America, [622].
• Biogenetic law, corollary of, [611].
• Birchall, E., local variation in British Lepidoptera, [60].
• Blanchard, on oviposition of Amblystoma, [594].
• Boisduval, figures of larvæ referred to, [216], [218], [245];
• on snake-like appearance of the larva of D. nicæa, [342];
• on the relationship of Euchloe belia and E. ausonia, [48].
• Bombycidæ, early stage of larvæ, [166].
• Bombyx. B. cynthia, colour changes in larva, [7];
• B. neustria, caterpillar refused by lizards, [336];
• B. Rubi, caterpillar eaten by lizards, [338].
• Boscher, E., dimorphism of larva of S. ocellatus, [241], [306].
• Brauer, insect phylogeny, [493].
• Burmeister, figures of larvæ referred to, [195], [196], [224], [232], [255], [261], [263], [436], [437];
• early stages of larvæ of Argentine butterflies, [166].
• Butler, A. G., affinities of genus Acronycta, [169];
• genera of Chærocampinæ, [190], [192], [194], [196];
• genera of Smerinthinæ, [233], [243], [244];
• genera of Macroglossinæ, [253], [254];
• species of Pterogon, [256];
• genera of Sphinginæ and Acherontiinæ, [262];
• on coloured larvæ being rejected, [336];
• affinities of Sesiidæ, [370];
• descriptions and figures of new Sphinx-larvæ, [524], [526].
• Bütschli, embryology of bees, [483].
• Butterflies, action of warmth in producing changes in colour and marking, [58];
• summer and winter larvæ and pupæ alike in seasonally dimorphic species, [64];
• seasonally dimorphic species hibernate as pupæ, [73];
• odoriferous organs, [102];
• early stages of larvæ, [165];
• di- and polymorphism of, [250];
• crepuscular habits of certain species, [475].
• C.
• Callosune eupompe, polymorphism of female, [251].
• Cameron, P., on larva of S. ocellatus, [241], [282];
• protective habits of saw-fly larvæ, [290], [294].
• Carlin, W. E., on Siredon lichenoides, [630].
• Caterpillars, origin of markings, [161];
• means of defence, [289];
• defensive habits, [290];
• food of brightly coloured species, [294];
• habit of concealment by day, [291], [296];
• phytophagic variability, [305], [531];
• seasonal variability, [305];
• experiments with ocellated species,
• 330;
• experiments with annulated species, [336];
• sexual dimorphism of, [308], [527], [534];
• independent variability of stages, [416].
• Cecidomyia, metagenesis of, [82], [586].
• Chærocampa. C. elpenor and C. porcellus, relationship of, [171];
• markings of larvæ of the genus, [177];
• C. elpenor, 1st and 2nd larval stages, [178];
• 3rd and 4th stages, [180];
• 5th stage, [181];
• 6th stage, [183];
• C. porcellus, 1st larval stage, [184];
• 2nd and 3rd stages, [185];
• 4th stage, [186];
• C. elpenor and porcellus, larval markings compared with other species of Chærocampa, [188];
• C. elpenor, experiments with larva, [331], [332];
• C. Japonica and C. Lewisii, larvæ, [194];
• C. cretica, larva, [524];
• C. lycetus, larva, [527];
• C. capensis, larva, [529];
• eaten by birds, &c., [530].
• Characters, new, backward transference of, [279].
• Chauvin, Fräulein von, experiments with Axolotl, [558];
• experiments with Salamandra atra, [615].
• Chavannes, figure of larva referred to, [255].
• Clarke, development of Amblystoma punctatum, [623].
• Classification, objects of, [168].
• Claus, on the origination of heterogenesis, [89].
• Clemens, larva of A. ello, [268].
• Climatic variation, [45];
• in P. Napi, [47];
• in E. Belia, [47];
• nature of causes producing, [52], [105];
• by sex, [61], [62];
• climatic varieties very distinct from local, [45];
• climatic influences operate slowly, [58];
• climatic changes gradual, [652], [660].
• Cole, B. G., seasonal dimorphism of E. punctaria, [4].
• Coleoptera, grub-like larvæ of, [495].
• Colias edusa, dimorphism of female, [250].
• Colour, biological value of, [289].
• Conditions of life, compulsory changes in, [651].
• Congruence in Lepidopterous families, [435];
• in Lepidopterous genera, [444].
• Coniferæ, food-plants of Anceryx, [268].
• Convergence of characters in Ophiuroidea, [396];
• in Dipterous larvæ, [494].
• Cope, Prof., habits of Siredon Mexicanus, [566];
• on distribution of Amblystoma, [569].
• Correlation of growth, [278], [363], [388], [415];
• action of, [415], [516];
• Von Hartmann’s views of, [670];
• nature of, [673].
• Cryptobranchus, characters of, [579].
• Cyclical propagation, [82], [699];
• and reversion, [613].
• D.
• Daphniacea, differences between allied species and genera, [516].
• Darwin, C., influence of external conditions of life, [59];
• inheritance at corresponding periods, [63], [275], [431];
• markings of butterflies, [101];
• definition of eye-spots, [326];
• adaptation in larvæ, [392];
• correlation of growth, [516];
• degeneration, [583];
• adaptation, [589];
• causes of sterility, [596];
• limited variability, [656];
• factors of variability, [676];
• abrupt variations, [701].
• Degeneration, [583];
• phyletic, [611].
• Deilephila, division of larvæ into groups, [199];
• D. Euphorbiæ, 1st larval stage, [202];
• 2nd and 3rd stages, [203];
• 4th and 5th stages, [205];
• D. nicæa, larva, [207];
• D. dahlii, larva, [208];
• D. vespertilio, larva, [209];
• D. Galii, larva, [211];
• 5th stage, [213];
• D. livornica, larva, [215];
• D. Zygophylli, larva, [217];
• D. Hippophaës, larva, [218], [650];
• division of larvæ according to phyletic stages, [223];
• genealogy of, [358];
• D. Galii, larva rejected, [340];
• D. Euphorbia, larva eaten, [340];
• habits of larvæ and imagines, [412];
• D. Robertsi, larva, [525].
• Development;
• of marking of Sphinx-larvæ, laws of, [274];
• phyletic, of markings of Sphingidæ, [370];
• unequal phyletic, [460].
• Dimorphism and polymorphism;
• of butterflies, [250];
• of Sphinx-larvæ, [300];
• of pupæ, [412].
• Diptera, incongruences among, [488];
• imaginal characters of, [488];
• divisions of, [498].
• Dodo, extinction of, [651].
• Dohrn, A.;
• on degeneration, [583];
• on functional change, [590].
• Dorfmeister, G.;
• experiments with A. levana, [11].
• Duponchel;
• figures of larvæ referred to, [207], [208], [218], [253].
• Duméril;
• experiments with Axolotl, [555], [564], [566], [593];
• characters of Axolotl, [375], [576].
• E.
• Edwards, W. H., seasonal dimorphism of P. pseudargiolus and P. violecea, [4];
• of P. Tharos and P. Marcia, [4];
• of P. Ajax, [30];
• of L. Artemis and L. Proserpina, [32];
• experiments with P. Ajax and P. Tharos, [33], [126], [140];
• pupal period of winter forms of P. Ajax, [36];
• experiments with G. interrogationis, [149];
• early stages of larvæ of N. American butterflies, [165];
• larva of P. pseudargiolus and attendant ants, [290];
• figures of larvæ referred to, [165], [436], [437].
• Eimer, colour variation in Lacerta muralis, [361].
• Emmelesia unifasciata, larva phytophagically variable, [307].
• Environment, direct and indirect action of, [686].
• Ephyra punctaria and E. omicronaria, seasonal dimorphism of, [4].
• Eriogaster lanestris, larva eaten by lizards, [336].
• Eriotus Pteridis, resemblance of larva, [320].
• Euchelia Jacobææ;
• larva and imago rejected by lizards, [336];
• constant in all three stages, [405].
• Euchloe:
• E. belia and E. ausonia, seasonal dimorphism of, [3];
• E. belia, climatic variation of, [47];
• var. Simplonia in Alps, [48];
• E. belemia, seasonal dimorphism and migration of, [78].
• Euproctus Rusconii, habits of, [617].
• Eusmerinthus Kindermanni, larva, [526].
• Evershed, J., larva of C. porcellus, [188].
• Eye-spots;
• repetition of, [277];
• development of, [325], [327];
• terrifying action, [330];
• resemble flower-buds, [334];
• useful when rudimentary, [366].
• F.
• Fatiot, Amphibia of Upper Engadine, [616].
• Fechner, on conscious matter, [714].
• Filippi, De, Triton alpestris, [585].
• Forel, Prof., fresh water Pulmonifera, [590].
• Form-relationship and blood-relationship, not always coincident, [395].
• Frantzius, Von, on climate of Mexico, [619].
• Frogs, [615], [616], [618];
• inedible species, [294].
• Function, change of, [365].
• G.
• Gegenbaur, Prof., the skull of Axolotl, [576].
• Gehrig, breeding of Axolotl, [565].
• Gené, habits of Euproctus Rusconii, [617].
• Genera, Lepidopterous, show greatest congruence, [459].
• Generation, heterogeneous, [679], [698], [702].
• Gentry, T. G., phytophagic variability of caterpillars, [306].
• Gerstäcker, division of the Diptera, [498].
• Geotriton fuscus, habits, [617].
• Glacial epoch; butterflies monogoneutic during, [19], [72];
• in N. America, [622].
• Gooch, W. D., description of larvæ referred to, [436];
• crepuscular butterflies, [476];
• sexual dimorphism of caterpillars, [535].
• Gooseberry, not increased in size, [654].
• Goss, H., larva of C. porcellus, [188].
• Gosse, P. H., embryonic larva of P. Homerus, [167].
• Grapta interrogationis, experiments with, [149].
• Green colour of caterpillars, [293], [310].
• Growth, law of, [655].
• Gynandromorphism in butterflies, [249].
• Gynanisa Isis, young larva, [166].
• H.
• Haeckel, Prof., on metagenesis, [80], [83];
• on ontogeny, [270];
• on adaptation, [281], [589];
• on the biogenetic law, [611];
• on reproduction and growth, [664];
• the plastidule theory, [667].
• Hare, white variation of, [660].
• Hartmann, E. von; his views examined, [645];
• on variability, [646], [652], [655], [664];
• on heredity, [657];
• his “Philosophy of the Unconscious,” [670];
• on correlation, [670], [673];
• on the nature of species, [671];
• on design in nature, [696];
• on the sensibility of atoms, [714].
• Helmholtz, Prof., on natural laws, [668];
• on the senses, [711].
• Hemaris hylas, larva, [255].
• Heredity, alternating and continuous, [24];
• cyclical, [66];
• homochronic, [63], [431];
• not mechanical, [657];
• nature of, [667].
• Herrich-Schäffer, wing neuration of butterflies, [449].
• Hertwig, on teeth of Amphibia, [376].
• Heterogenesis, definition of, [82], [96];
• parallelism between and metamorphosis, [86];
• origination of, [89].
• Heyden, Von, reproduction of Aphis, [97].
• Hilgendorf, on fossil shells, [109].
• Hipparchia semele, colour variations, [8].
• Hoffmann, Ernst, on immigration of European butterflies, [77].
• Horsfield and Moore, figures of caterpillars referred to, [193], [195], [196], [243], [253], [261].
• Hübner, figures of caterpillars referred to, [193], [208], [211], [218], [231], [267].
• Humboldt, Baron, on the Lake of Mexico, [608], [621].
• Huxley, Prof., “Anatomy of Invertebrated Animals” referred to, [498].
• Hydromedusæ, alternation of generations in, [83];
• want of parallelism in different generations of, [395].
• Hydrozoa, asexual reproduction and polymorphosis, [83].
• Hymenoptera, incongruence among, [481];
• imaginal characters, [481];
• causes of incongruence, [486].
• I.
• Ichthyodea, characters of, [577], [579].
• Incongruence; in Lepidopterous families, [437];
• genera, [446], [449];
• species, [451];
• varieties, [456];
• two kinds of, [458], [502];
• in Hymenoptera, [481];
• in Diptera, [488];
• different forms of, [503];
• dependent upon action of environment, [505], [507], [508].
• Inheritance; homochronic, [63], [431];
• definition of cyclical, [66];
• limited by sex, [68].
• Insects, ancestry of, [493].
• Intermaxillary gland of Axolotl, [623].
• J.
• Jullien, on Lissotriton punctatus, [591].
• K.
• Kant, referred to, [638], [709].
• Kirby, W. F., “Synonymic Catalogue” referred to, [2], [49], [435], [449].
• Kleemann, larva of S. Ligustri, [259].
• Knaggs, Dr. H. G., seasonal dimorphism of S. illustraria, [4].
• Kölliker, Prof., experiments with Axolotl, [564];
• characters of Axolotl, [575].
• L.
• Lacerta muralis, colour variations, [361].
• Lake of Mexico, [603], [608], [621].
• Lange, F. A., “History of Materialism” referred to, [645], [656].
• Langer, Prof., experiments with Pelobates tadpoles, [607].
• Lankester, Prof. E. R., on degeneration, [583].
• Lasiocampa Pini, larva eaten by lizards, [336];
• variable in two stages, [405].
• Lepidoptera; foes, [8];
• larva and imago independently variable, [401];
• species constant in three stages, [405];
• species variable in three stages, [405];
• variable in two stages, [405];
• variable in one stage, [406];
• causes of variability of pupæ, [411];
• case of incongruence among pupæ, [540].
• Leptodora hyalina, development, [93].
• Lepus timidus, white variations of, [662].
• Leuckart, reproduction of Ascaris nigrovenosa, [90];
• on growth, [654].
• Leydig, on sexual larvæ of Triton, [591].
• Limenitis Artemis and L. Proserpina, dimorphic in both sexes, [32];
• L. Camilla, constant in three stages, [405].
• Lines, dorsal, subdorsal, supra- and infra-spiracular, [175].
• Lissotriton punctatus, sexual larvæ, [591], [595].
• Lizards, colour variation of, [361];
• rejection of caterpillars by, [336].
• Local forms, distinct from climatic varieties, [45];
• produced by isolation, [109].
• Lockyer, B., young Noctua larvæ, [166].
• Lophostethus Dumolinii, larva, [527].
• Lophura hyas, larva, [254].
• Lubbock, Sir John, insect metamorphosis, [65], [431];
• derivation of metagenesis, [83];
• colours of caterpillars, [294];
• ancestry of insects, [493].
• Lycæna pseudargiolus, larva and ants, [290].
• M.
• Macroglossa, larvæ of, [245];
• M. stellatarum, oviposition of, [245];
• 1st, 2nd, and 3rd larval stages, [246];
• 4th and 5th stages, [247];
• di- and polymorphism of larva, [247];
• pupa of, [250];
• M. belis, and M. pyrrhosticta, larvæ, [255].
• Markings, general biological value of, [285];
• special biological value of, [308];
• subordinate, [347];
• of Sphingidæ, evolution of, [380].
• Marsh, Prof., on Siredon lichenoides, [567].
• Matter, assumption of, [711];
• conscious, [714].
• Mayer, Paul, insect phylogeny, [493].
• McLachlan, R., phytophagic variability, [305].
• Mechanism and teleology, [694].
• Medusæ, internal parasitism, [699].
• Melanopsis recurrens, from Paludina bed, [275].
• Menopoma, characters of, [579].
• Mérian, Madame, figures of caterpillars referred to, [195], [232], [263], [268].
• Metagenesis, defined, [81], [96];
• derivation, [83].
• Metamorphosis, [430].
• Mexico, climate of, [619].
• Millière, figure of larva referred to, [254].
• Mimicry, [648], [663].
• Möller, L., phytophagic variability, [305].
• Mono- and digoneutic species, [16].
• Moore, F., figures of caterpillars referred to, [436], [526].
• Morris, affinities of Apatura and Nymphalis, [438].
• Moths, seasonal dimorphism in, [4];
• resembling splinters, [292].
• Mühlenpfordt, on Lake of Mexico, [604].
• Müller, Fritz, odoriferous organs of butterflies, [102];
• sexual selection in butterflies, [103];
• habit and protective resemblance, [290];
• spiny caterpillars, [293];
• larva of Papilio Evander, [299];
• reversion in Orchids, [614];
• maggot-like larvæ of insects, [493];
• the biogenetic law, [611];
• mimicry, &c., [665].
• Müller, Hermann, larva of Stauropus Fagi, [290].
• Müller, P. E., Leptodora hyalina, [94].
• Murray, Andrew, seasonal change of colour, in animals, [7];
• green caterpillars, [293].
• Muscidæ, embryonic development, [492].
• N.
• Nägeli, reversion of cultivated plants, [107].
• Natural selection, [112], [361], [704], [705].
• Nature, mechanical conception of, [634];
• harmony of, [697].
• Neumayr and Paul, on Melanopsis, [275].
• Noctuæ, change of colour in larvæ, [166];
• number of legs in young larvæ, [166], [520];
• ontogeny of larvæ, [520].
• Noll, Dr., larva of Ach. Atropos, [324].
• North America, glaciation of, [622].
• Nurse-breeding in Hymenoptera and Diptera, [402].
• Nymphalidæ, larval classification, [435].
• O.
• Ontogeny, Fritz Müller and Haeckel on, [270];
• predominance of new characters in last stage of, [280], [283].
• Ontogeny and Phylogeny of Sphinx-markings, [177].
• Ophiuroidea, want of parallelism in, [396].
• Orchids, reversion in, [614].
• Organic compounds, synthesis of, [643].
• Organism, parts independently variable, [514].
• P.
• Pachylia Ficus, early stages of larva, [232].
• Packard, insect phylogeny, [493].
• Pangenesis, theory of, [668].
• Papilio. P. Ajax, seasonal forms, [30];
• P. Turnus, a dimorphic form, [32];
• P. Machaon and Podalirius, seasonally dimorphic in Spain and Italy, [74];
• P. Ajax, experiments with, [126];
• P. Merope, young larva, [166];
• P. Homerus, young larva, [167];
• P. Machaon, larva rejected by lizards, [339].
• Parallelism, phyletic, in metamorphic species, [390];
• phyletic, law of, [510].
• Pararga Ægeria, sexually dimorphic, [68];
• climatic variation of, [68].
• Pascoe, F. P., “Zoological Classification” referred to, [488], [498].
• Pelobates, experiments with tadpoles, [607].
• Pergesa Mongoliana, larva, [194].
• Philampelus achemon and P. satellitia, larvæ, [523];
• P. labruscæ, larva, [195];
• P. vitis, and P. anchemolus, larvæ, [232].
• Philosophy, function of, [640].
• Phyciodes Tharos and P. Marcia, seasonal dimorphism of, [4];
• experiments with, [140].
• Phylogeny, conclusions from, [270];
• and ontogeny of Sphinx-markings, [177];
• of insects, [493].
• Pierinæ, seasonal dimorphism of, [13];
• analogous seasonal dimorphism in, [60], [108];
• experiments with, [122].
• Pieris Napi, experiments with, [13];
• summer form the younger, [29];
• reversion of, caused by mechanical vibration, [38];
• var. Bryoniæ, the potential winter form, [39];
• experiments with this var., [40];
• this var. the parent form of Napi, [41];
• a climatic var., [41];
• very variable, [43];
• variability due to crossing, [43];
• var. æstiva more variable than var. vernalis, [43];
• P. Napi, summer and winter forms compared, [56];
• incongruence in this species, [457];
• P. Krueperi, seasonal dimorphism of, [78];
• P. Brassicæ, larva rejected by lizards, [337];
• constant in three stages, [405];
• photographic sensitiveness of pupa, [405];
• P. Napi, variable in two stages, [405].
• Plants, cultivated, [107];
• sterility of reverted, [596].
• Plastidule theory, [667].
• Plebeius amyntas and P. polysperchon, seasonal dimorphism, [2];
• P. Alexis, do., [3];
• P. pseudargiolus and P. violacea, do., [4];
• P. agestis, do., [50];
• polymorphism of females in this genus, [251].
• Pluvial period, [621].
• Polar animals, colour of, [658], [660].
• Polyommatus phlæas, distribution and climatic variation, [49];
• seasonal dimorphism, [50];
• Italian summer and winter forms compared, [55].
• Polyptychus dentatus, larva, [244].
• Ptarmigan, seasonal colours of, [6].
• Pterogon, larvæ of, [255];
• P. ænotheræ, larva, [256].
• Pulmonifera, functional change in lungs, [590].
• Pygæra bucephala, larva rejected by lizards, [337].
• Q.
• Quatrefages, De, spermatozoa of Amblystoma, [593].
• R.
• Rana temporaria of Upper Engadine, [618].
• Ratzeburg, larva of Anc. Pinastri, [265].
• Reproduction and growth, [664].
• Reversion, of species, [611];
• two modes of, [612];
• periodic, [613].
• Rhopalocera, characters of, [433], [471].
• Rhytina Stelleri, extinction of, [651].
• Riley, C. V., descriptions of caterpillars referred to, [437], [521], [522];
• on sexual dimorphism in caterpillar, [535].
• Ring-spots, [325];
• development, [327];
• signs of distastefulness, [341];
• resembling berries, [344].
• Rösel, breeding of Aras. Levana by, [6];
• on larva of Deil. Euphorbiæ, [206];
• of D. Galii, [213];
• of Smer. Tiliæ, [236];
• of S. ocellatus, [240];
• of Anc. Pinastri, [265], [268].
• Roux, Dr. W., struggle of parts in the organism, [689].
• Rutherford, D. G., colour variations of Ater. meleagris, [8].
• S.
• Sacc, on sterility of Amblystoma, [593].
• Salamanders, habits of Italian land species, [617];
• oviposition of, [630].
• Salamandra atra, experiments with, [615].
• Salamandrina, characters of, [577];
• S. perspicillata, habits of, [617].
• Sars, development of Leptodora hyalina, [94].
• Saturnia.
• S. Yamamai, variable in first stage only, [406];
• S. carpini, development of larva of local forms, [419].
• Satyrinæ, hibernation of larvæ, [73].
• Saussure, De, on Siredon Mexicanus, [565], [602];
• physical character of Mexican Lake, [603].
• Saw-flies, protective habits of larvæ, [290].
• Schmankewitsch, transformation of Artemia, [635].
• Schmidt, Oscar, on convergence of character, [396], [459];
• on variability, [654].
• Schreibers, sexual Triton larvæ, [591];
• arrested tadpoles, [607].
• Schulze, F. E., parasitism in Medusæ, [699].
• Science, function of, [640].
• Scudder, S. H., embryonic larvæ of butterflies, [166].
• Seasonal dimorphism, origin and significance, [1];
• seasonal change of colour in animals, [7];
• and climatic variation, [45];
• seasonal adaptation in caterpillars, [305].
• Seidlitz, reproduction of Axolotl, [571];
• white var. of hare, [662].
• Selenia tetralunaria, S. illunaria, S. lunaria, and S. illustraria, seasonal dimorphism, [4].
• Semper, C., origin of alternation of generations, [84];
• transformation of Amblystoma mavortium, [629].
• Semper, G., young Sphinx-larvæ, [166];
• figures of caterpillars referred to, [522].
• Senses, physiology of, [711].
• Sepp, figure of caterpillar referred to, [262].
• Sesiidæ, affinities of, [370].
• Sexual dimorphism, of butterflies, [32], [250];
• of caterpillars, [308], [527], [534];
• secondary sexual characters, [62];
• sexual selection, [69], [102].
• Siebold, Von, on Pulmonifera, [590].
• Siredon. S. Mexicanus, habits, [565];
• S. lichenoides, transformation, [567], [630];
• phyletic advance of genus, [584];
• retention of genus, [610];
• S. tigrinus from L. St. Isabel, [626].
• Slater, J. W., food of gaily coloured caterpillars, [294].
• Smerinthus, larvæ of, [232];
• S. Tiliæ, [233];
• 4th stage, [235];
• S. Populi, [236];
• and stage, [237];
• 3rd and 4th stages, [238];
• 5th stage, [239];
• S. ocellatus, [240];
• dimorphism of larva, [241];
• S. Tiliæ, variable in two stages, [405];
• North American species of, [453];
• S. tatarinovii, and S. planus, larvæ, [244].
• Species, causes of transformation of, [116];
• nature of, [671].
• Specific characters, [91];
• may originate through direct action, [100].
• Specific constitution, [112].
• Spencer, Herbert, theory of descent, [645];
• variability, [654];
• law of growth, [655].
• Spermatozoa, of Amblystoma, [593];
• of Lissotriton punctatus, [595].
• Speyer, Dr., seasonal dimorphism of moths, [4];
• of P. phlæas in Germany, [50].
• Sphingidæ, oviposition, [164];
• congruence and incongruence in genera, [450].
• Sphinx-markings, ontogeny and morphology, [177].
• Larvæ of genus, [259];
• S. Convolvuli, [650].
• Origination of markings, [273];
• law of development, [274];
• forms of markings, [309];
• S. Ligustri, constant in three stages, [405].
• Stainton, H. T., larval characters of Notodontidæ, [443];
• young larvæ of Triph. pronuba, [520].
• Staudinger, Dr., seasonal dimorphism of Euch. Belia and E. Ausonia, [3], [48];
• butterflies of Finnmark, [20];
• P. Napi var. Bryoniæ from Lapland, [44], [49];
• larvæ of Ch. alecto, [193];
• affinities of Nymphalis and Apatura, [438].
• Stauropus Fagi, protective habits of larva, [290].
• Sterility, causes of, [596];
• a result of reversion, [597];
• law of, in reversion forms, [599].
• Stoat, seasonal change of colour, [7].
• Strauch, “Revision of Salamandridæ” referred to, [570], [577].
• Strecker, H., N. American species of Smerinthus, [453].
• Stripes, longitudinal, [312], [372];
• oblique, [317], [373].
• T.
• Tadpoles, arrested development, [607].
• Tegetmeier, W. B., metamorphosis of Siredon, [566].
• Teleology and Mechanism, [694].
• Terebrantia, larvæ, [508].
• Transformation, causes of, [460], [496];
• factors of, [676].
• Trematoda, alternation of generations in, [83].
• Trimen, Roland;
• larva of Lopho. Dumolinii, [527];
• of Ch. Capensis, [529];
• phytophagic variability of larva of Ach. Atropos, [531].
• Triptogon roseipennis, larva, [244].
• Triton;
• T. alpestris, sexual larva, [585];
• reproductive larvæ, [590].
• Degeneration in genus, [612].
• Species of Upper Engadine, [631].
• Typical parts, modified by environment, [501], [513].
• U.
• Upper Engadine;
• Amphibia, [615];
• climate, [616];
• frog from, [618];
• Tritons of, [631].
• Urea, synthesis of, [643].
• V.
• Vanessa.
• V. Urticæ, climatic variation, [60];
• V. Atalanta, Urticæ, and polychloros, variable in two stages, [405];
• V. Io, variable in one stage, [407].
• Congruence and incongruence in genus, [446];
• phyletic completeness of, [447];
• arrangement of spines on larvæ, [448].
• Variability, origin, [107];
• limited, [114], [362], [653], [655];
• independent in different stages, [402];
• definition of, [404];
• a relative term, [408];
• causes of, in pupæ, [411];
• primary and secondary, [416];
• independent in different larval stages, [416];
• phytophagic, [305], [531];
• Von Hartmann’s views, [646];
• not unlimited, [647], [652], [655], [683];
• factors of, [676], [684];
• mechanical theory of, [677];
• individual, [681], [685].
• Variation; climatic, [45], [686];
• causes of, [105];
• analogous, [683];
• abrupt not inherited, [701].
• Varieties, climatic, distinct from local, [45].
• Velasco, Señor, Axolotl of L. St. Isabel, [626].
• Vital force, phyletic, elimination of, [287];
• objections to, [352], [461], [511], [518];
• ontogenetic, abandonment of, [641], [687].
• W.
• Wallace, A. R.;
• introduction of the expression “seasonal dimorphism,” [1];
• on the dull colours of female butterflies, [8];
• sexual dimorphism of butterflies, [32];
• local variation in colour, [60];
• comparison of seasonal dimorphism with alternation of generations, [80];
• the bright colours of caterpillars, [293];
• on adaptation, [589];
• on variability, [654], [658].
• Wallace, Dr., colour changes in larva of B. Cynthia, [7].
• Walsh, B. D., phytophagic variation, [305].
• Weale, J. P. M., on the young larvæ of Pap. merope and Gyn. Isis, [166];
• the larva of Ach. Atropos in S. Africa, [263];
• habits of S. African Sphinx-caterpillars, [290].
• Weir, J. Jenner, the colour variations of Hip. semele, [8];
• experiments with brightly-coloured, hairy, and spiny caterpillars, [336].
• Westwood, Prof. J. O., seasonal dimorphism of Eph. punctaria, [4];
• figure of caterpillar referred to, [243];
• gynandromorphic butterfly, [249].
• White colour, protection afforded by, [6], [659], [661].
• White, Dr. F. B., local variation in British Lepidoptera, [60];
• young Notua larvæ, [166].
• Wiedersheim, on the anatomy of Amblystoma, [577];
• habits of Geotriton fuscus, [617];
• anatomy of Axolotl, [623].
• Wigand, on the non-increase in size of the gooseberry, [654].
• Wilde, O., on larva of Dei. Galii, [213];
• of Dei. Hippophaës, [218];
• larval characters of the Notodontidæ, [443].
• Wilson, Owen, figure of larva of Ch. porcellus referred to, [188].
• Wöhler, Prof., the synthesis of urea, [643].
• Wood, T. W., photographic sensitiveness of Lepidopterous pupæ, [405].
• Würtemberger, on Ammonites, [275].
• Y.
• Yenisei, Swedish Expedition referred to, [20].
• Z.
• Zeller, P. C., seasonal dimorphism of Pl. Polysperchon and Pl. Amyntas, [2];
• of Italian butterflies, [75].

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