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that have since been urged against them. Moreover, the matter is by no means one of subordinate interest; it is the very kernel of the whole question of the reality and value of the principle of selection. For if selection alone does not suffice to explain “HARMONIOUS ADAPTATION” as I have called Spencer’s COADAPTATION, and if we require to call in the aid of the Lamarckian factor it would be questionable whether selection could explain any adaptations whatever. In this particular case–of worker bees –the Lamarckian factor may be excluded altogether, for it can be demonstrated that here at any rate the effects of use and disuse cannot be transmitted.

But if it be asked why we are unwilling to admit the cooperation of the Darwinian factor of selection and the Lamarckian factor, since this would afford us an easy and satisfactory explanation of the phenomena, I answer: BECAUSE THE LAMARCKIAN PRINCIPLE IS FALLACIOUS, AND BECAUSE BY ACCEPTING IT WE CLOSE THE WAY TOWARDS DEEPER INSIGHT. It is not a spirit of combativeness or a desire for self-vindication that induces me to take the field once more against the Lamarckian principle, it is the conviction that the progress of our knowledge is being obstructed by the acceptance of this fallacious principle, since the facile explanation it apparently affords prevents our seeking after a truer explanation and a deeper analysis.

The workers in the various species of ants are sterile, that is to say, they take no regular part in the reproduction of the species, although individuals among them may occasionally lay eggs. In addition to this they have lost the wings, and the receptaculum seminis, and their compound eyes have degenerated to a few facets. How could this last change have come about through disuse, since the eyes of workers are exposed to light in the same way as are those of the sexual insects and thus in this particular case are not liable to “disuse” at all? The same is true of the receptaculum seminis, which can only have been disused as far as its glandular portion and its stalk are concerned, and also of the wings, the nerves tracheae and epidermal cells of which could not cease to function until the whole wing had degenerated, for the chitinous skeleton of the wing does not function at all in the active sense.

But, on the other hand, the workers in all species have undergone modifications in a positive direction, as, for instance, the greater development of brain. In many species large workers have evolved,–the so- called SOLDIERS, with enormous jaws and teeth, which defend the colony,– and in others there are SMALL workers which have taken over other special functions, such as the rearing of the young Aphides. This kind of division of the workers into two castes occurs among several tropical species of ants, but it is also present in the Italian species, Colobopsis truncata. Beautifully as the size of the jaws could be explained as due to the increased use made of them by the “soldiers,” or the enlarged brain as due to the mental activities of the workers, the fact of the infertility of these forms is an insurmountable obstacle to accepting such an explanation. Neither jaws nor brain can have been evolved on the Lamarckian principle.

The problem of coadaptation is no easier in the case of the ant than in the case of the Giant Stag. Darwin himself gave a pretty illustration to show how imposing the difference between the two kinds of workers in one species would seem if we translated it into human terms. In regard to the Driver ants (Anomma) we must picture to ourselves a piece of work, “for instance the building of a house, being carried on by two kinds of workers, of which one group was five feet four inches high, the other sixteen feet high.” (“Origin of Species” (6th edition), page 232.)

Although the ant is a small animal as compared with man or with the Irish Elk, the “soldier” with its relatively enormous jaws is hardly less heavily burdened than the Elk with its antlers, and in the ant’s case, too, a strengthening of the skeleton, of the muscles, the nerves of the head, and of the legs must have taken place parallel with the enlargement of the jaws. HARMONIOUS ADAPTATION (coadaptation) has here been active in a high degree, and yet these “soldiers” are sterile! There thus remains nothing for it but to refer all their adaptations, positive and negative alike, to processes of selection which have taken place in the rudiments of the workers within the egg and sperm-cells of their parents. There is no way out of the difficulty except the one Darwin pointed out. He himself did not find the solution of the riddle at once. At first he believed that the case of the workers among social insects presented “the most serious special difficulty” in the way of his theory of natural selection; and it was only after it had become clear to him, that it was not the sterile insects themselves but their parents that were selected, according as they produced more or less well adapted workers, that he was able to refer to this very case of the conditions among ants “IN ORDER TO SHOW THE POWER OF NATURAL SELECTION” (“Origin of Species”, page 233; see also edition 1, page 242.). He explains his view by a simple but interesting illustration. Gardeners have produced, by means of long continued artificial selection, a variety of Stock, which bears entirely double, and therefore infertile flowers (Ibid. page 230.). Nevertheless the variety continues to be reproduced from seed, because in addition to the double and infertile flowers, the seeds always produce a certain number of single, fertile blossoms, and these are used to reproduce the double variety. These single and fertile plants correspond “to the males and females of an ant-colony, the infertile plants, which are regularly produced in large numbers, to the neuter workers of the colony.”

This illustration is entirely apt, the only difference between the two cases consisting in the fact that the variation in the flower is not a useful, but a disadvantageous one, which can only be preserved by artificial selection on the part of the gardener, while the transformations that have taken place parallel with the sterility of the ants are useful, since they procure for the colony an advantage in the struggle for existence, and they are therefore preserved by natural selection. Even the sterility itself in this case is not disadvantageous, since the fertility of the true females has at the same time considerably increased. We may therefore regard the sterile forms of ants, which have gradually been adapted in several directions to varying functions, AS A CERTAIN PROOF that selection really takes place in the germ-cells of the fathers and mothers of the workers, and that SPECIAL COMPLEXES OF PRIMORDIA (IDS) are present in the workers and in the males and females, and these complexes contain the primordia of the individual parts (DETERMINANTS). But since all living entities vary, the determinants must also vary, now in a favourable, now in an unfavourable direction. If a female produces eggs, which contain favourably varying determinants in the worker-ids, then these eggs will give rise to workers modified in the favourable direction, and if this happens with many females, the colony concerned will contain a better kind of worker than other colonies.

I digress here in order to give an account of the intimate processes, which, according to my view, take place within the germ-plasm, and which I have called “GERMINAL SELECTION.” These processes are of importance since they form the roots of variation, which in its turn is the root of natural selection. I cannot here do more than give a brief outline of the theory in order to show how the Darwin-Wallace theory of selection has gained support from it.

With others, I regard the minimal amount of substance which is contained within the nucleus of the germ-cells, in the form of rods, bands, or granules, as the GERM-SUBSTANCE or GERM-PLASM, and I call the individual granules IDS. There is always a multiplicity of such ids present in the nucleus, either occurring individually, or united in the form of rods or bands (chromosomes). Each id contains the primary constituents of a WHOLE individual, so that several ids are concerned in the development of a new individual.

In every being of complex structure thousands of primary constituents must go to make up a single id; these I call DETERMINANTS, and I mean by this name very small individual particles, far below the limits of microscopic visibility, vital units which feed, grow, and multiply by division. These determinants control the parts of the developing embryo,–in what manner need not here concern us. The determinants differ among themselves, those of a muscle are differently constituted from those of a nerve-cell or a glandular cell, etc., and every determinant is in its turn made up of minute vital units, which I call BIOPHORS, or the bearers of life. According to my view, these determinants not only assimilate, like every other living unit, but they VARY in the course of their growth, as every living unit does; they may vary qualitatively if the elements of which they are composed vary, they may grow and divide more or less rapidly, and their variations give rise to CORRESPONDING variations of the organ, cell, or cell-group which they determine. That they are undergoing ceaseless fluctuations in regard to size and quality seems to me the inevitable consequence of their unequal nutrition; for although the germ-cell as a whole usually receives sufficient nutriment, minute fluctuations in the amount carried to different parts within the germ-plasm cannot fail to occur.

Now, if a determinant, for instance of a sensory cell, receives for a considerable time more abundant nutriment than before, it will grow more rapidly–become bigger, and divide more quickly, and, later, when the id concerned develops into an embryo, this sensory cell will become stronger than in the parents, possibly even twice as strong. This is an instance of a HEREDITARY INDIVIDUAL VARIATION, arising from the germ.

The nutritive stream which, according to our hypothesis, favours the determinant N by chance, that is, for reasons unknown to us, may remain strong for a considerable time, or may decrease again; but even in the latter case it is conceivable that the ascending movement of the determinant may continue, because the strengthened determinant now ACTIVELY nourishes itself more abundantly,–that is to say, it attracts the nutriment to itself, and to a certain extent withdraws it from its fellow- determinants. In this way, it may–as it seems to me–get into PERMANENT UPWARD MOVEMENT, AND ATTAIN A DEGREE OF STRENGTH FROM WHICH THERE IS NO FALLING BACK. Then positive or negative selection sets in, favouring the variations which are advantageous, setting aside those which are disadvantageous.

In a similar manner a DOWNWARD variation of the determinants may take place, if its progress be started by a diminished flow of nutriment. The determinants which are weakened by this diminished flow will have less affinity for attracting nutriment because of their diminished strength, and they will assimilate more feebly and grow more slowly, unless chance streams of nutriment help them to recover themselves. But, as will presently be shown, a change of direction cannot take place at EVERY stage of the degenerative process. If a certain critical stage of downward progress be passed, even favourable conditions of food-supply will no longer suffice permanently to change the direction of the variation. Only two cases are conceivable; if the determinant corresponds to a USEFUL organ, only its removal can bring back the germ-plasm to its former level; therefore personal selection removes the id in question, with its determinants, from the germ-plasm, by causing the elimination of the individual in the struggle for existence. But there is another conceivable case; the determinants concerned may be those of an organ which has become USELESS, and they will then continue unobstructed, but with exceeding slowness, along the downward path, until the organ becomes vestigial, and finally disappears altogether.

The fluctuations of the determinants hither and thither may thus be transformed into a lasting ascending or descending movement; and THIS IS THE CRUCIAL POINT OF THESE GERMINAL PROCESSES.


The whole determinant system of an id, as I conceive it, is in a state of continual fluctuation upwards and downwards. In most cases the fluctuations will counteract one another, because the passive streams of nutriment soon change, but in many cases the limit from which a return is possible will be passed, and then the determinants concerned will continue to vary in the same direction, till they attain positive or negative selection-value. At this stage personal selection intervenes and sets aside the variation if it is disadvantageous, or favours–that is to say, preserves–it if it is advantageous. Only THE DETERMINANT OF A USELESS ORGAN IS UNINFLUENCED BY PERSONAL SELECTION, and, as experience shows, it sinks downwards; that is, the organ that corresponds to it degenerates very slowly but uninterruptedly till, after what must obviously be an immense stretch of time, it disappears from the germ-plasm altogether.

Thus we find in the fact of the degeneration of disused parts the proof that not all the fluctuations of a determinant return to equilibrium again, but that, when the movement has attained to a certain strength, it continues IN THE SAME DIRECTION. We have entire certainty in regard to this as far as the downward progress is concerned, and we must assume it also in regard to ascending variations, as the phenomena of artificial selection certainly justify us in doing. If the Japanese breeders were able to lengthen the tail feathers of the cock to six feet, it can only have been because the determinants of the tail-feathers in the germ-plasm had already struck out a path of ascending variation, and this movement was taken advantage of by the breeder, who continually selected for reproduction the individuals in which the ascending variation was most marked. For all breeding depends upon the unconscious selection of germinal variations.

Of course these germinal processes cannot be proved mathematically, since we cannot actually see the play of forces of the passive fluctuations and their causes. We cannot say how great these fluctuations are, and how quickly or slowly, how regularly or irregularly they change. Nor do we know how far a determinant must be strengthened by the passive flow of the nutritive stream if it is to be beyond the danger of unfavourable variations, or how far it must be weakened passively before it loses the power of recovering itself by its own strength. It is no more possible to bring forward actual proofs in this case than it was in regard to the selection-value of the initial stages of an adaptation. But if we consider that all heritable variations must have their roots in the germ-plasm, and further, that when personal selection does not intervene, that is to say, in the case of parts which have become useless, a degeneration of the part, and therefore also of its determinant must inevitably take place; then we must conclude that processes such as I have assumed are running their course within the germ-plasm, and we can do this with as much certainty as we were able to infer, from the phenomena of adaptation, the selection- value of their initial stages. The fact of the degeneration of disused parts seems to me to afford irrefutable proof that the fluctuations within the germ-plasm ARE THE REAL ROOT OF ALL HEREDITARY VARIATION, and the preliminary condition for the occurrence of the Darwin-Wallace factor of selection. Germinal selection supplies the stones out of which personal selection builds her temples and palaces: ADAPTATIONS. The importance for the theory of the process of degeneration of disused parts cannot be over- estimated, especially when it occurs in sterile animal forms, where we are free from the doubt as to the alleged LAMARCKIAN FACTOR which is apt to confuse our ideas in regard to other cases.

If we regard the variation of the many determinants concerned in the transformation of the female into the sterile worker as having come about through the gradual transformation of the ids into worker-ids, we shall see that the germ-plasm of the sexual ants must contain three kinds of ids, male, female, and worker ids, or if the workers have diverged into soldiers and nest-builders, then four kinds. We understand that the worker-ids arose because their determinants struck out a useful path of variation, whether upward or downward, and that they continued in this path until the highest attainable degree of utility of the parts determined was reached. But in addition to the organs of positive or negative selection-value, there were some which were indifferent as far as the success and especially the functional capacity of the workers was concerned: wings, ovarian tubes, receptaculum seminis, a number of the facets of the eye, perhaps even the whole eye. As to the ovarian tubes it is possible that their degeneration was an advantage for the workers, in saving energy, and if so selection would favour the degeneration; but how could the presence of eyes diminish the usefulness of the workers to the colony? or the minute receptaculum seminis, or even the wings? These parts have therefore degenerated BECAUSE THEY WERE OF NO FURTHER VALUE TO THE INSECT. But if selection did not influence the setting aside of these parts because they were neither of advantage nor of disadvantage to the species, then the Darwinian factor of selection is here confronted with a puzzle which it cannot solve alone, but which at once becomes clear when germinal selection is added. For the determinants of organs that have no further value for the organism, must, as we have already explained, embark on a gradual course of retrograde development.

In ants the degeneration has gone so far that there are no wing-rudiments present in ANY species, as is the case with so many butterflies, flies, and locusts, but in the larvae the imaginal discs of the wings are still laid down. With regard to the ovaries, degeneration has reached different levels in different species of ants, as has been shown by the researches of my former pupil, Elizabeth Bickford. In many species there are twelve ovarian tubes, and they decrease from that number to one; indeed, in one species no ovarian tube at all is present. So much at least is certain from what has been said, that in this case EVERYTHING depends on the fluctuations of the elements of the germ-plasm. Germinal selection, here as elsewhere, presents the variations of the determinants, and personal selection favours or rejects these, or,–if it be a question of organs which have become useless,–it does not come into play at all, and allows the descending variation free course.

It is obvious that even the problem of COADAPTATION IN STERILE ANIMALS can thus be satisfactorily explained. If the determinants are oscillating upwards and downwards in continual fluctuation, and varying more pronouncedly now in one direction now in the other, useful variations of every determinant will continually present themselves anew, and may, in the course of generations, be combined with one another in various ways. But there is one character of the determinants that greatly facilitates this complex process of selection, that, after a certain limit has been reached, they go on varying in the same direction. From this it follows that development along a path once struck out may proceed without the continual intervention of personal selection. This factor only operates, so to speak, at the beginning, when it selects the determinants which are varying in the right direction, and again at the end, when it is necessary to put a check upon further variation. In addition to this, enormously long periods have been available for all these adaptations, as the very gradual transition stages between females and workers in many species plainly show, and thus this process of transformation loses the marvellous and mysterious character that seemed at the first glance to invest it, and takes rank, without any straining, among the other processes of selection. It seems to me that, from the facts that sterile animal forms can adapt themselves to new vital functions, their superfluous parts degenerate, and the parts more used adapt themselves in an ascending direction, those less used in a descending direction, we must draw the conclusion that harmonious adaptation here comes about WITHOUT THE COOPERATION OF THE LAMARCKIAN PRINCIPLE. This conclusion once established, however, we have no reason to refer the thousands of cases of harmonious adaptation, which occur in exactly the same way among other animals or plants, to a principle, the ACTIVE INTERVENTION OF WHICH IN THE TRANSFORMATION OF SPECIES IS NOWHERE PROVED. WE DO NOT REQUIRE IT TO EXPLAIN THE FACTS, AND THEREFORE WE MUST NOT ASSUME IT.

The fact of coadaptation, which was supposed to furnish the strongest argument against the principle of selection, in reality yields the clearest evidence in favour of it. We MUST assume it, BECAUSE NO OTHER POSSIBILITY OF EXPLANATION IS OPEN TO US, AND BECAUSE THESE ADAPTATIONS ACTUALLY EXIST, THAT IS TO SAY, HAVE REALLY TAKEN PLACE. With this conviction I attempted, as far back as 1894, when the idea of germinal selection had not yet occurred to me, to make “harmonious adaptation” (coadaptation) more easily intelligible in some way or other, and so I was led to the idea, which was subsequently expounded in detail by Baldwin, and Lloyd Morgan, and also by Osborn, and Gulick as ORGANIC SELECTION. It seemed to me that it was not necessary that all the germinal variations required for secondary variations should have occurred SIMULTANEOUSLY, since, for instance, in the case of the stag, the bones, muscles, sinews, and nerves would be incited by the increasing heaviness of the antlers to greater activity in THE INDIVIDUAL LIFE, and so would be strengthened. The antlers can only have increased in size by very slow degrees, so that the muscles and bones may have been able to keep pace with their growth in the individual life, until the requisite germinal variations presented themselves. In this way a disharmony between the increasing weight of the antlers and the parts which support and move them would be avoided, since time would be given for the appropriate germinal variations to occur, and so to set agoing the HEREDITARY variation of the muscles, sinews, and bones. (“The Effect of External Influences upon Development”, Romanes Lecture, Oxford, 1894.)

I still regard this idea as correct, but I attribute less importance to “organic selection” than I did at that time, in so far that I do not believe that it ALONE could effect complex harmonious adaptations. Germinal selection now seems to me to play the chief part in bringing about such adaptations. Something the same is true of the principle I have called “Panmixia”. As I became more and more convinced, in the course of years, that the LAMARCKIAN PRINCIPLE ought not to be called in to explain the dwindling of disused parts, I believed that this process might be simply explained as due to the cessation of the conservative effect of natural selection. I said to myself that, from the moment in which a part ceases to be of use, natural selection withdraws its hand from it, and then it must inevitably fall from the height of its adaptiveness, because inferior variants would have as good a chance of persisting as better ones, since all grades of fitness of the part in question would be mingled with one another indiscriminately. This is undoubtedly true, as Romanes pointed out ten years before I did, and this mingling of the bad with the good probably does bring about a deterioration of the part concerned. But it cannot account for the steady diminution, which always occurs when a part is in process of becoming rudimentary, and which goes on until it ultimately disappears altogether. The process of dwindling cannot therefore be explained as due to panmixia alone; we can only find a sufficient explanation in germinal selection.


The impetus in all directions given by Darwin through his theory of selection has been an immeasurable one, and its influence is still felt. It falls within the province of the historian of science to enumerate all the ideas which, in the last quarter of the nineteenth century, grew out of Darwin’s theories, in the endeavour to penetrate more deeply into the problem of the evolution of the organic world. Within the narrow limits to which this paper is restricted, I cannot attempt to discuss any of these.



Sexual selection goes hand in hand with natural selection. From the very first I have regarded sexual selection as affording an extremely important and interesting corroboration of natural selection, but, singularly enough, it is precisely against this theory that an adverse judgment has been pronounced in so many quarters, and it is only quite recently, and probably in proportion as the wealth of facts in proof of it penetrates into a wider circle, that we seem to be approaching a more general recognition of this side of the problem of adaptation. Thus Darwin’s words in his preface to the second edition (1874) of his book, “The Descent of Man and Sexual Selection”, are being justified: “My conviction as to the operation of natural selection remains unshaken,” and further, “If naturalists were to become more familiar with the idea of sexual selection, it would, I think, be accepted to a much greater extent, and already it is fully and favourably accepted by many competent judges.” Darwin was able to speak thus because he was already acquainted with an immense mass of facts, which, taken together, yield overwhelming evidence of the validity of the principle of sexual selection.

NATURAL SELECTION chooses out for reproduction the individuals that are best equipped for the struggle for existence, and it does so at every stage of development; it thus improves the species in all its stages and forms. SEXUAL SELECTION operates only on individuals that are already capable of reproduction, and does so only in relation to the attainment of reproduction. It arises from the rivalry of one sex, usually the male, for the possession of the other, usually the female. Its influence can therefore only DIRECTLY affect one sex, in that it equips it better for attaining possession of the other. But the effect may extend indirectly to the female sex, and thus the whole species may be modified, without, however, becoming any more capable of resistance in the struggle for existence, for sexual selection only gives rise to adaptations which are likely to give their possessor the victory over rivals in the struggle for possession of the female, and which are therefore peculiar to the wooing sex: the manifold “secondary sexual characters.” The diversity of these characters is so great that I cannot here attempt to give anything approaching a complete treatment of them, but I should like to give a sufficient number of examples to make the principle itself, in its various modes of expression, quite clear.

One of the chief preliminary postulates of sexual selection is the unequal number of individuals in the two sexes, for if every male immediately finds his mate there can be no competition for the possession of the female. Darwin has shown that, for the most part, the inequality between the sexes is due simply to the fact that there are more males than females, and therefore the males must take some pains to secure a mate. But the inequality does not always depend on the numerical preponderance of the males, it is often due to polygamy; for, if one male claims several females, the number of females in proportion to the rest of the males will be reduced. Since it is almost always the males that are the wooers, we must expect to find the occurrence of secondary sexual characters chiefly among them, and to find it especially frequent in polygamous species. And this is actually the case.

If we were to try to guess–without knowing the facts–what means the male animals make use of to overcome their rivals in the struggle for the possession of the female, we might name many kinds of means, but it would be difficult to suggest any which is not actually employed in some animal group or other. I begin with the mere difference in strength, through which the male of many animals is so sharply distinguished from the female, as, for instance, the lion, walrus, “sea-elephant,” and others. Among these the males fight violently for the possession of the female, who falls to the victor in the combat. In this simple case no one can doubt the operation of selection, and there is just as little room for doubt as to the selection-value of the initial stages of the variation. Differences in bodily strength are apparent even among human beings, although in their case the struggle for the possession of the female is no longer decided by bodily strength alone.

Combats between male animals are often violent and obstinate, and the employment of the natural weapons of the species in this way has led to perfecting of these, e.g. the tusks of the boar, the antlers of the stag, and the enormous, antler-like jaws of the stag-beetle. Here again it is impossible to doubt that variations in these organs presented themselves, and that these were considerable enough to be decisive in combat, and so to lead to the improvement of the weapon.

Among many animals, however, the females at first withdraw from the males; they are coy, and have to be sought out, and sometimes held by force. This tracking and grasping of the females by the males has given rise to many different characters in the latter, as, for instance, the larger eyes of the male bee, and especially of the males of the Ephemerids (May-flies), some species of which show, in addition to the usual compound eyes, large, so-called turban-eyes, so that the whole head is covered with seeing surfaces. In these species the females are very greatly in the minority (1- 100), and it is easy to understand that a keen competition for them must take place, and that, when the insects of both sexes are floating freely in the air, an unusually wide range of vision will carry with it a decided advantage. Here again the actual adaptations are in accordance with the preliminary postulates of the theory. We do not know the stages through which the eye has passed to its present perfected state, but, since the number of simple eyes (facets) has become very much greater in the male than in the female, we may assume that their increase is due to a gradual duplication of the determinants of the ommatidium in the germ-plasm, as I have already indicated in regard to sense-organs in general. In this case, again, the selection-value of the initial stages hardly admits of doubt; better vision DIRECTLY secures reproduction.

In many cases THE ORGAN OF SMELL shows a similar improvement. Many lower Crustaceans (Daphnidae) have better developed organs of smell in the male sex. The difference is often slight and amounts only to one or two olfactory filaments, but certain species show a difference of nearly a hundred of these filaments (Leptodora). The same thing occurs among insects.

We must briefly consider the clasping or grasping organs which have developed in the males among many lower Crustaceans, but here natural selection plays its part along with sexual selection, for the union of the sexes is an indispensable condition for the maintenance of the species, and as Darwin himself pointed out, in many cases the two forms of selection merge into each other. This fact has always seemed to me to be a proof of natural selection, for, in regard to sexual selection, it is quite obvious that the victory of the best-equipped could have brought about the improvement only of the organs concerned, the factors in the struggle, such as the eye and the olfactory organ.

We come now to the EXCITANTS; that is, to the group of sexual characters whose origin through processes of selection has been most frequently called in question. We may cite the LOVE-CALLS produced by many male insects, such as crickets and cicadas. These could only have arisen in animal groups in which the female did not rapidly flee from the male, but was inclined to accept his wooing from the first. Thus, notes like the chirping of the male cricket serve to entice the females. At first they were merely the signal which showed the presence of a male in the neighbourhood, and the female was gradually enticed nearer and nearer by the continued chirping. The male that could make himself heard to the greatest distance would obtain the largest following, and would transmit the beginnings, and, later, the improvement of his voice to the greatest number of descendants. But sexual excitement in the female became associated with the hearing of the love-call, and then the sound-producing organ of the male began to improve, until it attained to the emission of the long-drawn-out soft notes of the mole-cricket or the maenad-like cry of the cicadas. I cannot here follow the process of development in detail, but will call attention to the fact that the original purpose of the voice, the announcing of the male’s presence, became subsidiary, and the exciting of the female became the chief goal to be aimed at. The loudest singers awakened the strongest excitement, and the improvement resulted as a matter of course. I conceive of the origin of bird-song in a somewhat similar manner, first as a means of enticing, then of exciting the female.

One more kind of secondary sexual character must here be mentioned: the odour which emanates from so many animals at the breeding season. It is possible that this odour also served at first merely to give notice of the presence of individuals of the other sex, but it soon became an excitant, and as the individuals which caused the greatest degree of excitement were preferred, it reached as high a pitch of perfection as was possible to it. I shall confine myself here to the comparatively recently discovered fragrance of butterflies. Since Fritz Muller found out that certain Brazilian butterflies gave off fragrance “like a flower,” we have become acquainted with many such cases, and we now know that in all lands, not only many diurnal Lepidoptera but nocturnal ones also give off a delicate odour, which is agreeable even to man. The ethereal oil to which this fragrance is due is secreted by the skin-cells, usually of the wing, as I showed soon after the discovery of the SCENT-SCALES. This is the case in the males; the females have no SPECIAL scent-scales recognisable as such by their form, but they must, nevertheless, give off an extremely delicate fragrance, although our imperfect organ of smell cannot perceive it, for the males become aware of the presence of a female, even at night, from a long distance off, and gather round her. We may therefore conclude, that both sexes have long given forth a very delicate perfume, which announced their presence to others of the same species, and that in many species (NOT IN ALL) these small beginnings became, in the males, particularly strong scent-scales of characteristic form (lute, brush, or lyre-shaped). At first these scales were scattered over the surface of the wing, but gradually they concentrated themselves, and formed broad, velvety bands, or strong, prominent brushes, and they attained their highest pitch of evolution when they became enclosed within pits or folds of the skin, which could be opened to let the delicious fragrance stream forth suddenly towards the female. Thus in this case also we see that characters, the original use of which was to bring the sexes together, and so to maintain the species, have been evolved in the males into means for exciting the female. And we can hardly doubt, that the females are most readily enticed to yield to the butterfly that sends out the strongest fragrance,–that is to say, that excites them to the highest degree. It is a pity that our organs of smell are not fine enough to examine the fragrance of male Lepidoptera in general, and to compare it with other perfumes which attract these insects. (See Poulton, “Essays on Evolution”, 1908, pages 316, 317.) As far as we can perceive them they resemble the fragrance of flowers, but there are Lepidoptera whose scent suggests musk. A smell of musk is also given off by several plants: it is a sexual excitant in the musk-deer, the musk-sheep, and the crocodile.

As far as we know, then, it is perfumes similar to those of flowers that the male Lepidoptera give off in order to entice their mates, and this is a further indication that animals, like plants, can to a large extent meet the claims made upon them by life, and produce the adaptations which are most purposive,–a further proof, too, of my proposition that the useful variations, so to speak, are ALWAYS THERE. The flowers developed the perfumes which entice their visitors, and the male Lepidoptera developed the perfumes which entice and excite their mates.

There are many pretty little problems to be solved in this connection, for there are insects, such as some flies, that are attracted by smells which are unpleasant to us, like those from decaying flesh and carrion. But there are also certain flowers, some orchids for instance, which give forth no very agreeable odour, but one which is to us repulsive and disgusting; and we should therefore expect that the males of such insects would give off a smell unpleasant to us, but there is no case known to me in which this has been demonstrated.

In cases such as we have discussed, it is obvious that there is no possible explanation except through selection. This brings us to the last kind of secondary sexual characters, and the one in regard to which doubt has been most frequently expressed,–decorative colours and decorative forms, the brilliant plumage of the male pheasant, the humming-birds, and the bird of Paradise, as well as the bright colours of many species of butterfly, from the beautiful blue of our little Lycaenidae to the magnificent azure of the large Morphinae of Brazil. In a great many cases, though not by any means in all, the male butterflies are “more beautiful” than the females, and in the Tropics in particular they shine and glow in the most superb colours. I really see no reason why we should doubt the power of sexual selection, and I myself stand wholly on Darwin’s side. Even though we certainly cannot assume that the females exercise a conscious choice of the “handsomest” mate, and deliberate like the judges in a court of justice over the perfections of their wooers, we have no reason to doubt that distinctive forms (decorative feathers) and colours have a particularly exciting effect upon the female, just as certain odours have among animals of so many different groups, including the butterflies. The doubts which existed for a considerable time, as a result of fallacious experiments, as to whether the colours of flowers really had any influence in attracting butterflies have now been set at rest through a series of more careful investigations; we now know that the colours of flowers are there on account of the butterflies, as Sprengel first showed, and that the blossoms of Phanerogams are selected in relation to them, as Darwin pointed out.

Certainly it is not possible to bring forward any convincing proof of the origin of decorative colours through sexual selection, but there are many weighty arguments in favour of it, and these form a body of presumptive evidence so strong that it almost amounts to certainty.

In the first place, there is the analogy with other secondary sexual characters. If the song of birds and the chirping of the cricket have been evolved through sexual selection, if the penetrating odours of male animals,–the crocodile, the musk-deer, the beaver, the carnivores, and, finally, the flower-like fragrances of the butterflies have been evolved to their present pitch in this way, why should decorative colours have arisen in some other way? Why should the eye be less sensitive to SPECIFICALLY MALE colours and other VISIBLE signs ENTICING TO THE FEMALE, than the olfactory sense to specifically male odours, or the sense of hearing to specifically male sounds? Moreover, the decorative feathers of birds are almost always spread out and displayed before the female during courtship. I have elsewhere (“The Evolution Theory”, London, 1904, I. page 219.) pointed out that decorative colouring and sweet-scentedness may replace one another in Lepidoptera as well as in flowers, for just as some modestly coloured flowers (mignonette and violet) have often a strong perfume, while strikingly coloured ones are sometimes quite devoid of fragrance, so we find that the most beautiful and gaily-coloured of our native Lepidoptera, the species of Vanessa, have no scent-scales, while these are often markedly developed in grey nocturnal Lepidoptera. Both attractions may, however, be combined in butterflies, just as in flowers. Of course, we cannot explain why both means of attraction should exist in one genus, and only one of them in another, since we do not know the minutest details of the conditions of life of the genera concerned. But from the sporadic distribution of scent-scales in Lepidoptera, and from their occurrence or absence in nearly related species, we may conclude that fragrance is a relatively MODERN acquirement, more recent than brilliant colouring.

One thing in particular that stamps decorative colouring as a product of selection is ITS GRADUAL INTENSIFICATION by the addition of new spots, which we can quite well observe, because in many cases the colours have been first acquired by the males, and later transmitted to the females by inheritance. The scent-scales are never thus transmitted, probably for the same reason that the decorative colours of many birds are often not transmitted to the females: because with these they would be exposed to too great elimination by enemies. Wallace was the first to point out that in species with concealed nests the beautiful feathers of the male occurred in the female also, as in the parrots, for instance, but this is not the case in species which brood on an exposed nest. In the parrots one can often observe that the general brilliant colouring of the male is found in the female, but that certain spots of colour are absent, and these have probably been acquired comparatively recently by the male and have not yet been transmitted to the female.

Isolation of the group of individuals which is in process of varying is undoubtedly of great value in sexual selection, for even a solitary conspicuous variation will become dominant much sooner in a small isolated colony, than among a large number of members of a species.

Anyone who agrees with me in deriving variations from germinal selection will regard that process as an essential aid towards explaining the selection of distinctive courtship-characters, such as coloured spots, decorative feathers, horny outgrowths in birds and reptiles, combs, feather-tufts, and the like, since the beginnings of these would be presented with relative frequency in the struggle between the determinants within the germ-plasm. The process of transmission of decorative feathers to the female results, as Darwin pointed out and illustrated by interesting examples, in the COLOUR-TRANSFORMATION OF A WHOLE SPECIES, and this process, as the phyletically older colouring of young birds shows, must, in the course of thousands of years, have repeated itself several times in a line of descent.

If we survey the wealth of phenomena presented to us by secondary sexual characters, we can hardly fail to be convinced of the truth of the principle of sexual selection. And certainly no one who has accepted natural selection should reject sexual selection, for, not only do the two processes rest upon the same basis, but they merge into one another, so that it is often impossible to say how much of a particular character depends on one and how much on the other form of selection.


An actual proof of the theory of sexual selection is out of the question, if only because we cannot tell when a variation attains to selection-value. It is certain that a delicate sense of smell is of value to the male moth in his search for the female, but whether the possession of one additional olfactory hair, or of ten, or of twenty additional hairs leads to the success of its possessor we are unable to tell. And we are groping even more in the dark when we discuss the excitement caused in the female by agreeable perfumes, or by striking and beautiful colours. That these do make an impression is beyond doubt; but we can only assume that slight intensifications of them give any advantage, and we MUST assume this SINCE OTHERWISE SECONDARY SEXUAL CHARACTERS REMAIN INEXPLICABLE.

The same thing is true in regard to natural selection. It is not possible to bring forward any actual proof of the selection-value of the initial stages, and the stages in the increase of variations, as has been already shown. But the selection-value of a finished adaptation can in many cases be statistically determined. Cesnola and Poulton have made valuable experiments in this direction. The former attached forty-five individuals of the green, and sixty-five of the brown variety of the praying mantis (Mantis religiosa), by a silk thread to plants, and watched them for seventeen days. The insects which were on a surface of a colour similar to their own remained uneaten, while twenty-five green insects on brown parts of plants had all disappeared in eleven days.

The experiments of Poulton and Sanders (“Report of the British Association” (Bristol, 1898), London, 1899, pages 906-909.) were made with 600 pupae of Vanessa urticae, the “tortoise-shell butterfly.” The pupae were artificially attached to nettles, tree-trunks, fences, walls, and to the ground, some at Oxford, some at St Helens in the Isle of Wight. In the course of a month 93 per cent of the pupae at Oxford were killed, chiefly by small birds, while at St Helens 68 per cent perished. The experiments showed very clearly that the colour and character of the surface on which the pupa rests–and thus its own conspicuousness–are of the greatest importance. At Oxford only the four pupae which were fastened to nettles emerged; all the rest–on bark, stones and the like–perished. At St Helens the elimination was as follows: on fences where the pupae were conspicuous, 92 per cent; on bark, 66 per cent; on walls, 54 per cent; and among nettles, 57 per cent. These interesting experiments confirm our views as to protective coloration, and show further, THAT THE RATIO OF ELIMINATION IN THE SPECIES IS A VERY HIGH ONE, AND THAT THEREFORE SELECTION MUST BE VERY KEEN.

We may say that the process of selection follows as a logical necessity from the fulfilment of the three preliminary postulates of the theory: variability, heredity, and the struggle for existence, with its enormous ratio of elimination in all species. To this we must add a fourth factor, the INTENSIFICATION of variations which Darwin established as a fact, and which we are now able to account for theoretically on the basis of germinal selection. It may be objected that there is considerable uncertainty about this LOGICAL proof, because of our inability to demonstrate the selection- value of the initial stages and the individual stages of increase. We have therefore to fall back on PRESUMPTIVE EVIDENCE. This is to be found in THE INTERPRETATIVE VALUE OF THE THEORY. Let us consider this point in greater detail.

In the first place, it is necessary to emphasise what is often overlooked, namely, that the theory not only explains the TRANSFORMATIONS of species, it also explains THEIR REMAINING THE SAME; in addition to the principle of varying, it contains within itself that of PERSISTING. It is part of the essence of selection, that it not only causes a part to VARY till it has reached its highest pitch of adaptation, but that it MAINTAINS IT AT THIS PITCH. THIS CONSERVING INFLUENCE OF NATURAL SELECTION is of great importance, and was early recognised by Darwin; it follows naturally from the principle of the survival of the fittest.

We understand from this how it is that a species which has become fully adapted to certain conditions of life ceases to vary, but remains “constant,” as long as the conditions of life FOR IT remain unchanged, whether this be for thousands of years, or for whole geological epochs. But the most convincing proof of the power of the principle of selection lies in the innumerable multitude of phenomena which cannot be explained in any other way. To this category belong all structures which are only PASSIVELY of advantage to the organism, because none of these can have arisen by the alleged LAMARCKIAN PRINCIPLE. These have been so often discussed that we need do no more than indicate them here. Until quite recently the sympathetic coloration of animals–for instance, the whiteness of Arctic animals–was referred, at least in part, to the DIRECT influence of external factors, but the facts can best be explained by referring them to the processes of selection, for then it is unnecessary to make the gratuitous assumption that many species are sensitive to the stimulus of cold and that others are not. The great majority of Arctic land-animals, mammals and birds, are white, and this proves that they were all able to present the variation which was most useful for them. The sable is brown, but it lives in trees, where the brown colouring protects and conceals it more effectively. The musk-sheep (Ovibos moschatus) is also brown, and contrasts sharply with the ice and snow, but it is protected from beasts of prey by its gregarious habit, and therefore it is of advantage to be visible from as great a distance as possible. That so many species have been able to give rise to white varieties does not depend on a special sensitiveness of the skin to the influence of cold, but to the fact that Mammals and Birds have a general tendency to vary towards white. Even with us, many birds–starlings, blackbirds, swallows, etc.–occasionally produce white individuals, but the white variety does not persist, because it readily falls a victim to the carnivores. This is true of white fawns, foxes, deer, etc. The whiteness, therefore, arises from internal causes, and only persists when it is useful. A great many animals living in a GREEN ENVIRONMENT have become clothed in green, especially insects, caterpillars, and Mantidae, both persecuted and persecutors.

That it is not the direct effect of the environment which calls forth the green colour is shown by the many kinds of caterpillar which rest on leaves and feed on them, but are nevertheless brown. These feed by night and betake themselves through the day to the trunk of the tree, and hide in the furrows of the bark. We cannot, however, conclude from this that they were UNABLE to vary towards green, for there are Arctic animals which are white only in winter and brown in summer (Alpine hare, and the ptarmigan of the Alps), and there are also green leaf-insects which remain green only while they are young and difficult to see on the leaf, but which become brown again in the last stage of larval life, when they have outgrown the leaf. They then conceal themselves by day, sometimes only among withered leaves on the ground, sometimes in the earth itself. It is interesting that in one genus, Chaerocampa, one species is brown in the last stage of larval life, another becomes brown earlier, and in many species the last stage is not wholly brown, a part remaining green. Whether this is a case of a double adaptation, or whether the green is being gradually crowded out by the brown, the fact remains that the same species, even the same individual, can exhibit both variations. The case is the same with many of the leaf-like Orthoptera, as, for instance, the praying mantis (Mantis religiosa) which we have already mentioned.

But the best proofs are furnished by those often-cited cases in which the insect bears a deceptive resemblance to another object. We now know many such cases, such as the numerous imitations of green or withered leaves, which are brought about in the most diverse ways, sometimes by mere variations in the form of the insect and in its colour, sometimes by an elaborate marking, like that which occurs in the Indian leaf-butterflies, Kallima inachis. In the single butterfly-genus Anaea, in the woods of South America, there are about a hundred species which are all gaily coloured on the upper surface, and on the reverse side exhibit the most delicate imitation of the colouring and pattern of a leaf, generally without any indication of the leaf-ribs, but extremely deceptive nevertheless. Anyone who has seen only one such butterfly may doubt whether many of the insignificant details of the marking can really be of advantage to the insect. Such details are for instance the apparent holes and splits in the apparently dry or half-rotten leaf, which are usually due to the fact that the scales are absent on a circular or oval patch so that the colourless wing-membrane lies bare, and one can look through the spot as through a window. Whether the bird which is seeking or pursuing the butterflies takes these holes for dewdrops, or for the work of a devouring insect, does not affect the question; the mirror-like spot undoubtedly increases the general deceptiveness, for the same thing occurs in many leaf-butterflies, though not in all, and in some cases it is replaced in quite a peculiar manner. In one species of Anaea (A. divina), the resting butterfly looks exactly like a leaf out of the outer edge of which a large semicircular piece has been eaten, possibly by a caterpillar; but if we look more closely it is obvious that there is no part of the wing absent, and that the semicircular piece is of a clear, pale yellow colour, while the rest of the wing is of a strongly contrasted dark brown.

But the deceptive resemblance may be caused in quite a different manner. I have often speculated as to what advantage the brilliant white C could give to the otherwise dusky-coloured “Comma butterfly” (Grapta C. album). Poulton’s recent observations (“Proc. Ent. Soc”., London, May 6, 1903.) have shown that this represents the imitation of a crack such as is often seen in dry leaves, and is very conspicuous because the light shines through it.

The utility obviously lies in presenting to the bird the very familiar picture of a broken leaf with a clear shining slit, and we may conclude, from the imitation of such small details, that the birds are very sharp observers and that the smallest deviation from the usual arrests their attention and incites them to closer investigation. It is obvious that such detailed–we might almost say such subtle–deceptive resemblances could only have come about in the course of long ages through the acquirement from time to time of something new which heightened the already existing resemblance.

In face of facts like these there can be no question of chance, and no one has succeeded so far in finding any other explanation to replace that by selection. For the rest, the apparent leaves are by no means perfect copies of a leaf; many of them only represent the torn or broken piece, or the half or two-thirds of a leaf, but then the leaves themselves frequently do not present themselves to the eye as a whole, but partially concealed among other leaves. Even those butterflies which, like the species of Kallima and Anaea, represent the whole of a leaf with stalk, ribs, apex, and the whole breadth, are not actual copies which would satisfy a botanist; there is often much wanting. In Kallima the lateral ribs of the leaf are never all included in the markings; there are only two or three on the left side and at most four or five on the right, and in many individuals these are rather obscure, while in others they are comparatively distinct. This furnishes us with fresh evidence in favour of their origin through processes of selection, for a botanically perfect picture could not arise in this way; there could only be a fixing of such details as heightened the deceptive resemblance.

Our postulate of origin through selection also enables us to understand why the leaf-imitation is on the lower surface of the wing in the diurnal Lepidoptera, and on the upper surface in the nocturnal forms, corresponding to the attitude of the wings in the resting position of the two groups.

The strongest of all proofs of the theory, however, is afforded by cases of true “mimicry,” those adaptations discovered by Bates in 1861, consisting in the imitation of one species by another, which becomes more and more like its model. The model is always a species that enjoys some special protection from enemies, whether because it is unpleasant to taste, or because it is in some way dangerous.

It is chiefly among insects and especially among butterflies that we find the greatest number of such cases. Several of these have been minutely studied, and every detail has been investigated, so that it is difficult to understand how there can still be disbelief in regard to them. If the many and exact observations which have been carefully collected and critically discussed, for instance by Poulton (“Essays on Evolution”, 1889-1907, Oxford, 1908, passim, e.g. page 269.) were thoroughly studied, the arguments which are still frequently urged against mimicry would be found untenable; we can hardly hope to find more convincing proof of the actuality of the processes of selection than these cases put into our hands. The preliminary postulates of the theory of mimicry have been disputed, for instance, that diurnal butterflies are persecuted and eaten by birds, but observations specially directed towards this point in India, Africa, America and Europe have placed it beyond all doubt. If it were necessary I could myself furnish an account of my own observations on this point.

In the same way it has been established by experiment and observation in the field that in all the great regions of distribution there are butterflies which are rejected by birds and lizards, their chief enemies, on account of their unpleasant smell or taste. These butterflies are usually gaily and conspicuously coloured and thus–as Wallace first interpreted it–are furnished with an easily recognisable sign: a sign of unpalatableness or WARNING COLOURS. If they were not thus recognisable easily and from a distance, they would frequently be pecked at by birds, and then rejected because of their unpleasant taste; but as it is, the insect-eaters recognise them at once as unpalatable booty and ignore them. Such IMMUNE (The expression does not refer to all the enemies of this butterfly; against ichneumon-flies, for instance, their unpleasant smell usually gives no protection.) species, wherever they occur, are imitated by other palatable species, which thus acquire a certain degree of protection.

It is true that this explanation of the bright, conspicuous colours is only a hypothesis, but its foundations,–unpalatableness, and the liability of other butterflies to be eaten,–are certain, and its consequences–the existence of mimetic palatable forms–confirm it in the most convincing manner. Of the many cases now known I select one, which is especially remarkable, and which has been thoroughly investigated, Papilio dardanus (merope), a large, beautiful, diurnal butterfly which ranges from Abyssinia throughout the whole of Africa to the south coast of Cape Colony.

The males of this form are everywhere ALMOST the same in colour and in form of wings, save for a few variations in the sparse black markings on the pale yellow ground. But the females occur in several quite different forms and colourings, and one of these only, the Abyssinian form, is like the male, while the other three or four are MIMETIC, that is to say, they copy a butterfly of quite a different family the Danaids, which are among the IMMUNE forms. In each region the females have thus copied two or three different immune species. There is much that is interesting to be said in regard to these species, but it would be out of keeping with the general tenor of this paper to give details of this very complicated case of polymorphism in P. dardanus. Anyone who is interested in the matter will find a full and exact statement of the case in as far as we know it, in Poulton’s “Essays on Evolution” (pages 373-375). (Professor Poulton has corrected some wrong descriptions which I had unfortunately overlooked in the Plates of my book “Vortrage uber Descendenztheorie”, and which refer to Papilio dardanus (merope). These mistakes are of no importance as far as and understanding of the mimicry-theory is concerned, but I hope shortly to be able to correct them in a later edition.) I need only add that three different mimetic female forms have been reared from the eggs of a single female in South Africa. The resemblance of these forms to their immune models goes so far that even the details of the LOCAL forms of the models are copied by the mimetic species.

It remains to be said that in Madagascar a butterfly, Papilio meriones, occurs, of which both sexes are very similar in form and markings to the non-mimetic male of P. dardanus, so that it probably represents the ancestor of this latter species.

In face of such facts as these every attempt at another explanation must fail. Similarly all the other details of the case fulfil the preliminary postulates of selection, and leave no room for any other interpretation. That the males do not take on the protective colouring is easily explained, because they are in general more numerous, and the females are more important for the preservation of the species, and must also live longer in order to deposit their eggs. We find the same state of things in many other species, and in one case (Elymnias undularis) in which the male is also mimetically coloured, it copies quite a differently coloured immune species from the model followed by the female. This is quite intelligible when we consider that if there were TOO MANY false immune types, the birds would soon discover that there were palatable individuals among those with unpalatable warning colours. Hence the imitation of different immune species by Papilio dardanus!

I regret that lack of space prevents my bringing forward more examples of mimicry and discussing them fully. But from the case of Papilio dardanus alone there is much to be learnt which is of the highest importance for our understanding of transformations. It shows us chiefly what I once called, somewhat strongly perhaps, THE OMNIPOTENCE OF NATURAL SELECTION in answer to an opponent who had spoken of its “inadequacy.” We here see that one and the same species is capable of producing four or five different patterns of colouring and marking; thus the colouring and marking are not, as has often been supposed, a necessary outcome of the specific nature of the species, but a true adaptation, which cannot arise as a direct effect of climatic conditions, but solely through what I may call the sorting out of the variations produced by the species, according to their utility. That caterpillars may be either green or brown is already something more than could have been expected according to the old conception of species, but that one and the same butterfly should be now pale yellow, with black; now red with black and pure white; now deep black with large, pure white spots; and again black with a large ochreous-yellow spot, and many small white and yellow spots; that in one sub-species it may be tailed like the ancestral form, and in another tailless like its Danaid model,–all this shows a far-reaching capacity for variation and adaptation that wide never have expected if we did not see the facts before us. How it is possible that the primary colour-variations should thus be intensified and combined remains a puzzle even now; we are reminded of the modern three-colour printing,–perhaps similar combinations of the primary colours take place in this case; in any case the direction of these primary variations is determined by the artist whom we know as natural selection, for there is no other conceivable way in which the model could affect the butterfly that is becoming more and more like it. The same climate surrounds all four forms of female; they are subject to the same conditions of nutrition. Moreover, Papilio dardanus is by no means the only species of butterfly which exhibits different kinds of colour-pattern on its wings. Many species of the Asiatic genus Elymnias have on the upper surface a very good imitation of an immune Euploeine (Danainae), often with a steel-blue ground-colour, while the under surface is well concealed when the butterfly is at rest,– thus there are two kinds of protective coloration each with a different meaning! The same thing may be observed in many non-mimetic butterflies, for instance in all our species of Vanessa, in which the under side shows a grey-brown or brownish-black protective coloration, but we do not yet know with certainty what may be the biological significance of the gaily coloured upper surface.

In general it may be said that mimetic butterflies are comparatively rare species, but there are exceptions, for instance Limenitis archippus in North America, of which the immune model (Danaida plexippus) also occurs in enormous numbers.

In another mimicry-category the imitators are often more numerous than the models, namely in the case of the imitation of DANGEROUS INSECTS by harmless species. Bees and wasps are dreaded for their sting, and they are copied by harmless flies of the genera Eristalis and Syrphus, and these mimics often occur in swarms about flowering plants without damage to themselves or to their models; they are feared and are therefore left unmolested.

In regard also to the FAITHFULNESS OF THE COPY the facts are quite in harmony with the theory, according to which the resemblance must have arisen and increased BY DEGREES. We can recognise this in many cases, for even now the mimetic species show very VARYING DEGREES OF RESEMBLANCE to their immune model. If we compare, for instance, the many different imitators of Danaida chrysippus we find that, with their brownish-yellow ground-colour, and the position and size, and more or less sharp limitation of their clear marginal spots, they have reached very different degrees of nearness to their model. Or compare the female of Elymnias undularis with its model Danaida genutia; there is a general resemblance, but the marking of the Danaida is very roughly imitated in Elymnias.

Another fact that bears out the theory of mimicry is, that even when the resemblance in colour-pattern is very great, the WING-VENATION, which is so constant, and so important in determining the systematic position of butterflies, is never affected by the variation. The pursuers of the butterfly have no time to trouble about entomological intricacies.

I must not pass over a discovery of Poulton’s which is of great theoretical importance–that mimetic butterflies may reach the same effect by very different means. (“Journ. Linn. Soc. London (Zool.)”, Vol. XXVI. 1898, pages 598-602.) Thus the glass-like transparency of the wing of a certain Ithomiine (Methona) and its Pierine mimic (Dismorphia orise) depends on a diminution in the size of the scales; in the Danaine genus Ituna it is due to the fewness of the scales, and in a third imitator, a moth (Castnia linus var. heliconoides) the glass-like appearance of the wing is due neither to diminution nor to absence of scales, but to their absolute colourlessness and transparency, and to the fact that they stand upright. In another moth mimic (Anthomyza) the arrangement of the transparent scales is normal. Thus it is not some unknown external influence that has brought about the transparency of the wing in these five forms, as has sometimes been supposed. Nor is it a hypothetical INTERNAL evolutionary tendency, for all three vary in a different manner. The cause of this agreement can only lie in selection, which preserves and intensifies in each species the favourable variations that present themselves. The great faithfulness of the copy is astonishing in these cases, for it is not THE WHOLE wing which is transparent; certain markings are black in colour, and these contrast sharply with the glass-like ground. It is obvious that the pursuers of these butterflies must be very sharp-sighted, for otherwise the agreement between the species could never have been pushed so far. The less the enemies see and observe, the more defective must the imitation be, and if they had been blind, no visible resemblance between the species which required protection could ever have arisen.

A seemingly irreconcilable contradiction to the mimicry theory is presented in the following cases, which were known to Bates, who, however, never succeeded in bringing them into line with the principle of mimicry.

In South America there are, as we have already said, many mimics of the immune Ithomiinae (or as Bates called them Heliconidae). Among these there occur not merely species which are edible, and thus require the protection of a disguise, but others which are rejected on account of their unpalatableness. How could the Ithomiine dress have developed in their case, and of what use is it, since the species would in any case be immune? In Eastern Brazil, for instance, there are four butterflies, which bear a most confusing resemblance to one another in colour, marking, and form of wing, and all four are unpalatable to birds. They belong to four different genera and three sub-families, and we have to inquire: Whence came this resemblance and what end does it serve? For a long time no satisfactory answer could be found, but Fritz Muller (In “Kosmos”, 1879, page 100.), seventeen years after Bates, offered a solution to the riddle, when he pointed out that young birds could not have an instinctive knowledge of the unpalatableness of the Ithomiines, but must learn by experience which species were edible and which inedible. Thus each young bird must have tasted at least one individual of each inedible species and discovered its unpalatability, before it learnt to avoid, and thus to spare the species. But if the four species resemble each other very closely the bird will regard them all as of the same kind, and avoid them all. Thus there developed a process of selection which resulted in the survival of the Ithomiine-like individuals, and in so great an increase of resemblance between the four species, that they are difficult to distinguish one from another even in a collection. The advantage for the four species, living side by side as they do e.g. in Bahia, lies in the fact that only one individual from the MIMICRY-RING (“inedible association”) need be tasted by a young bird, instead of at least four individuals, as would otherwise be the case. As the number of young birds is great, this makes a considerable difference in the ratio of elimination.

These interesting mimicry-rings (trusts), which have much significance for the theory, have been the subject of numerous and careful investigations, and at least their essential features are now fully established. Muller took for granted, without making any investigations, that young birds only learn by experience to distinguish between different kinds of victims. But Lloyd Morgan’s (“Habit and Instinct”, London, 1896.) experiments with young birds proved that this is really the case, and at the same time furnished an additional argument against the LAMARCKIAN PRINCIPLE.

In addition to the mimicry-rings first observed in South America, others have been described from Tropical India by Moore, and by Poulton and Dixey from Africa, and we may expect to learn many more interesting facts in this connection. Here again the preliminary postulates of the theory are satisfied. And how much more that would lead to the same conclusion might be added!

As in the case of mimicry many species have come to resemble one another through processes of selection, so we know whole classes of phenomena in which plants and animals have become adapted to one another, and have thus been modified to a considerable degree. I refer particularly to the relation between flowers and insects; but as there is an article on “The Biology of Flowers” in this volume, I need not discuss the subject, but will confine myself to pointing out the significance of these remarkable cases for the theory of selection. Darwin has shown that the originally inconspicuous blossoms of the phanerogams were transformed into flowers through the visits of insects, and that, conversely, several large orders of insects have been gradually modified by their association with flowers, especially as regards the parts of their body actively concerned. Bees and butterflies in particular have become what they are through their relation to flowers. In this case again all that is apparently contradictory to the theory can, on closer investigation, be beautifully interpreted in corroboration of it. Selection can give rise only to what is of use to the organism actually concerned, never to what is of use to some other organism, and we must therefore expect to find that in flowers only characters of use to THEMSELVES have arisen, never characters which are of use to insects only, and conversely that in the insects characters useful to them and not merely to the plants would have originated. For a long time it seemed as if an exception to this rule existed in the case of the fertilisation of the yucca blossoms by a little moth, Pronuba yuccasella. This little moth has a sickle-shaped appendage to its mouth-parts which occurs in no other Lepidopteron, and which is used for pushing the yellow pollen into the opening of the pistil, thus fertilising the flower. Thus it appears as if a new structure, which is useful only to the plant, has arisen in the insect. But the difficulty is solved as soon as we learn that the moth lays its eggs in the fruit-buds of the Yucca, and that the larvae, when they emerge, feed on the developing seeds. In effecting the fertilisation of the flower the moth is at the same time making provision for its own offspring, since it is only after fertilisation that the seeds begin to develop. There is thus nothing to prevent our referring this structural adaptation in Pronuba yuccasella to processes of selection, which have gradually transformed the maxillary palps of the female into the sickle-shaped instrument for collecting the pollen, and which have at the same time developed in the insect the instinct to press the pollen into the pistil.

In this domain, then, the theory of selection finds nothing but corroboration, and it would be impossible to substitute for it any other explanation, which, now that the facts are so well known, could be regarded as a serious rival to it. That selection is a factor, and a very powerful factor in the evolution of organisms, can no longer be doubted. Even although we cannot bring forward formal proofs of it IN DETAIL, cannot calculate definitely the size of the variations which present themselves, and their selection-value, cannot, in short, reduce the whole process to a mathematical formula, yet we must assume selection, because it is the only possible explanation applicable to whole classes of phenomena, and because, on the other hand, it is made up of factors which we know can be proved actually to exist, and which, IF they exist, must of logical necessity cooperate in the manner required by the theory. WE MUST ACCEPT IT BECAUSE THE PHENOMENA OF EVOLUTION AND ADAPTATION MUST HAVE A NATURAL BASIS, AND BECAUSE IT IS THE ONLY POSSIBLE EXPLANATION OF THEM. (This has been discussed in many of my earlier works. See for instance “The All- Sufficiency of Natural Selection, a reply to Herbert Spencer”, London, 1893.)

Many people are willing to admit that selection explains adaptations, but they maintain that only a part of the phenomena are thus explained, because everything does not depend upon adaptation. They regard adaptation as, so to speak, a special effort on the part of Nature, which she keeps in readiness to meet particularly difficult claims of the external world on organisms. But if we look at the matter more carefully we shall find that adaptations are by no means exceptional, but that they are present everywhere in such enormous numbers, that it would be difficult in regard to any structure whatever, to prove that adaptation had NOT played a part in its evolution.

How often has the senseless objection been urged against selection that it can create nothing, it can only reject. It is true that it cannot create either the living substance or the variations of it; both must be given. But in rejecting one thing it preserves another, intensifies it, combines it, and in this way CREATES what is new. EVERYTHING in organisms depends on adaptation; that is to say, everything must be admitted through the narrow door of selection, otherwise it can take no part in the building up of the whole. But, it is asked, what of the direct effect of external conditions, temperature, nutrition, climate and the like? Undoubtedly these can give rise to variations, but they too must pass through the door of selection, and if they cannot do this they are rejected, eliminated from the constitution of the species.

It may, perhaps, be objected that such external influences are often of a compelling power, and that every animal MUST submit to them, and that thus selection has no choice and can neither select nor reject. There may be such cases; let us assume for instance that the effect of the cold of the Arctic regions was to make all the mammals become black; the result would be that they would all be eliminated by selection, and that no mammals would be able to live there at all. But in most cases a certain percentage of animals resists these strong influences, and thus selection secures a foothold on which to work, eliminating the unfavourable variation, and establishing a useful colouring, consistent with what is required for the maintenance of the species.

Everything depends upon adaptation! We have spoken much of adaptation in colouring, in connection with the examples brought into prominence by Darwin, because these are conspicuous, easily verified, and at the same time convincing for the theory of selection. But is it only desert and polar animals whose colouring is determined through adaptation? Or the leaf-butterflies, and the mimetic species, or the terrifying markings, and “warning-colours” and a thousand other kinds of sympathetic colouring? It is, indeed, never the colouring alone which makes up the adaptation; the structure of the animal plays a part, often a very essential part, in the protective disguise, and thus MANY variations may cooperate towards ONE common end. And it is to be noted that it is by no means only external parts that are changed; internal parts are ALWAYS modified at the same time–for instance, the delicate elements of the nervous system on which depend the INSTINCT of the insect to hold its wings, when at rest, in a perfectly definite position, which, in the leaf-butterfly, has the effect of bringing the two pieces on which the marking occurs on the anterior and posterior wing into the same direction, and thus displaying as a whole the fine curve of the midrib on the seeming leaf. But the wing-holding instinct is not regulated in the same way in all leaf-butterflies; even our indigenous species of Vanessa, with their protective ground-colouring, have quite a distinctive way of holding their wings so that the greater part of the anterior wing is covered by the posterior when the butterfly is at rest. But the protective colouring appears on the posterior wing and on the tip of the anterior, TO PRECISELY THE DISTANCE TO WHICH IT IS LEFT UNCOVERED. This occurs, as Standfuss has shown, in different degree in our two most nearly allied species, the uncovered portion being smaller in V. urticae than in V. polychloros. In this case, as in most leaf-butterflies, the holding of the wing was probably the primary character; only after that was thoroughly established did the protective marking develop. In any case, the instinctive manner of holding the wings is associated with the protective colouring, and must remain as it is if the latter is to be effective. How greatly instincts may change, that is to say, may be adapted, is shown by the case of the Noctuid “shark” moth, Xylina vetusta. This form bears a most deceptive resemblance to a piece of rotten wood, and the appearance is greatly increased by the modification of the innate impulse to flight common to so many animals, which has here been transformed into an almost contrary instinct. This moth does not fly away from danger, but “feigns death,” that is, it draws antennae, legs and wings close to the body, and remains perfectly motionless. It may be touched, picked up, and thrown down again, and still it does not move. This remarkable instinct must surely have developed simultaneously with the wood-colouring; at all events, both cooperating variations are now present, and prove that both the external and the most minute internal structure have undergone a process of adaptation.

The case is the same with all structural variations of animal parts, which are not absolutely insignificant. When the insects acquired wings they must also have acquired the mechanism with which to move them–the musculature, and the nervous apparatus necessary for its automatic regulation. All instincts depend upon compound reflex mechanisms and are just as indispensable as the parts they have to set in motion, and all may have arisen through processes of selection if the reasons which I have elsewhere given for this view are correct. (“The Evolution Theory”, London, 1904, page 144.)

Thus there is no lack of adaptations within the organism, and particularly in its most important and complicated parts, so that we may say that there is no actively functional organ that has not undergone a process of adaptation relative to its function and the requirements of the organism. Not only is every gland structurally adapted, down to the very minutest histological details, to its function, but the function is equally minutely adapted to the needs of the body. Every cell in the mucous lining of the intestine is exactly regulated in its relation to the different nutritive substances, and behaves in quite a different way towards the fats, and towards nitrogenous substances, or peptones.

I have elsewhere called attention to the many adaptations of the whale to the surrounding medium, and have pointed out–what has long been known, but is not universally admitted, even now–that in it a great number of important organs have been transformed in adaptation to the peculiar conditions of aquatic life, although the ancestors of the whale must have lived, like other hair-covered mammals, on land. I cited a number of these transformations–the fish-like form of the body, the hairlessness of the skin, the transformation of the fore-limbs to fins, the disappearance of the hind-limbs and the development of a tail fin, the layer of blubber under the skin, which affords the protection from cold necessary to a warm- blooded animal, the disappearance of the ear-muscles and the auditory passages, the displacement of the external nares to the forehead for the greater security of the breathing-hole during the brief appearance at the surface, and certain remarkable changes in the respiratory and circulatory organs which enable the animal to remain for a long time under water. I might have added many more, for the list of adaptations in the whale to aquatic life is by no means exhausted; they are found in the histological structure and in the minutest combinations in the nervous system. For it is obvious that a tail-fin must be used in quite a different way from a tail, which serves as a fly-brush in hoofed animals, or as an aid to springing in the kangaroo or as a climbing organ; it will require quite different reflex-mechanisms and nerve-combinations in the motor centres.

I used this example in order to show how unnecessary it is to assume a special internal evolutionary power for the phylogenesis of species, for this whole order of whales is, so to speak, MADE UP OF ADAPTATIONS; it deviates in many essential respects from the usual mammalian type, and all the deviations are adaptations to aquatic life. But if precisely the most essential features of the organisation thus depend upon adaptation, what is left for a phyletic force to do, since it is these essential features of the structure it would have to determine? There are few people now who believe in a phyletic evolutionary power, which is not made up of the forces known to us–adaptation and heredity–but the conviction that EVERY part of an organism depends upon adaptation has not yet gained a firm footing. Nevertheless, I must continue to regard this conception as the correct one, as I have long done.

I may be permitted one more example. The feather of a bird is a marvellous structure, and no one will deny that as a whole it depends upon adaptation. But what part of it DOES NOT depend upon adaptation? The hollow quill, the shaft with its hard, thin, light cortex, and the spongy substance within it, its square section compared with the round section of the quill, the flat barbs, their short, hooked barbules which, in the flight-feathers, hook into one another with just sufficient firmness to resist the pressure of the air at each wing-beat, the lightness and firmness of the whole apparatus, the elasticity of the vane, and so on. And yet all this belongs to an organ which is only passively functional, and therefore can have nothing to do with the LAMARCKIAN PRINCIPLE. Nor can the feather have arisen through some magical effect of temperature, moisture, electricity, or specific nutrition, and thus selection is again our only anchor of safety.

But–it will be objected–the substance of which the feather consists, this peculiar kind of horny substance, did not first arise through selection in the course of the evolution of the birds, for it formed the covering of the scales of their reptilian ancestors. It is quite true that a similar substance covered the scales of the Reptiles, but why should it not have arisen among them through selection? Or in what other way could it have arisen, since scales are also passively useful parts? It is true that if we are only to call adaptation what has been acquired by the species we happen to be considering, there would remain a great deal that could not be referred to selection; but we are postulating an evolution which has stretched back through aeons, and in the course of which innumerable adaptations took place, which had not merely ephemeral persistence in a genus, a family or a class, but which was continued into whole Phyla of animals, with continual fresh adaptations to the special conditions of each species, family, or class, yet with persistence of the fundamental elements. Thus the feather, once acquired, persisted in all birds, and the vertebral column, once gained by adaptation in the lowest forms, has persisted in all the Vertebrates, from Amphioxus upwards, although with constant readaptation to the conditions of each particular group. Thus everything we can see in animals is adaptation, whether of to-day, or of yesterday, or of ages long gone by; every kind of cell, whether glandular, muscular, nervous, epidermic, or skeletal, is adapted to absolutely definite and specific functions, and every organ which is composed of these different kinds of cells contains them in the proper proportions, and in the particular arrangement which best serves the function of the organ; it is thus adapted to its function.


But all adaptations CAN be referred to selection; the only point that remains doubtful is whether they all MUST be referred to it.

However that may be, whether the LAMARCKIAN PRINCIPLE is a factor that has cooperated with selection in evolution, or whether it is altogether fallacious, the fact remains, that selection is the cause of a great part of the phyletic evolution of organisms on our earth. Those who agree with me in rejecting the LAMARCKIAN PRINCIPLE will regard selection as the only GUIDING factor in evolution, which creates what is new out of the transmissible variations, by ordering and arranging these, selecting them in relation to their number and size, as the architect does his building- stones so that a particular style must result. (“Variation under Domestication”, 1875 II. pages 426, 427.) But the building-stones themselves, the variations, have their basis in the influences which cause variation in those vital units which are handed on from one generation to another, whether, taken together they form the WHOLE organism, as in Bacteria and other low forms of life, or only a germ-substance, as in unicellular and multicellular organisms. (The Author and Editor are indebted to Professor Poulton for kindly assisting in the revision of the proof of this Essay.)


Professor of Botany in the University of Amsterdam.


Before Darwin, little was known concerning the phenomena of variability. The fact, that hardly two leaves on a tree were exactly the same, could not escape observation: small deviations of the same kind were met with everywhere, among individuals as well as among the organs of the same plant. Larger aberrations, spoken of as monstrosities, were for a long time regarded as lying outside the range of ordinary phenomena. A special branch of inquiry, that of Teratology, was devoted to them, but it constituted a science by itself, sometimes connected with morphology, but having scarcely any bearing on the processes of evolution and heredity.

Darwin was the first to take a broad survey of the whole range of variations in the animal and vegetable kingdoms. His theory of Natural Selection is based on the fact of variability. In order that this foundation should be as strong as possible he collected all the facts, scattered in the literature of his time, and tried to arrange them in a scientific way. He succeeded in showing that variations may be grouped along a line of almost continuous gradations, beginning with simple differences in size and ending with monstrosities. He was struck by the fact that, as a rule, the smaller the deviations, the more frequently they appear, very abrupt breaks in characters being of rare occurrence.

Among these numerous degrees of variability Darwin was always on the look out for those which might, with the greatest probability, be considered as affording material for natural selection to act upon in the development of new species. Neither of the extremes complied with his conceptions. He often pointed out, that there are a good many small fluctuations, which in this respect must be absolutely useless. On the other hand, he strongly combated the belief, that great changes would be necessary to explain the origin of species. Some authors had propounded the idea that highly adapted organs, e.g. the wings of a bird, could not have been developed in any other way than by a comparatively sudden modification of a well defined and important kind. Such a conception would allow of great breaks or discontinuity in the evolution of highly differentiated animals and plants, shortening the time for the evolution of the whole organic kingdom and getting over numerous difficulties inherent in the theory of slow and gradual progress. It would, moreover, account for the genetic relation of the larger groups of both animals and plants. It would, in a word, undoubtedly afford an easy means of simplifying the problem of descent with modification.

Darwin, however, considered such hypotheses as hardly belonging to the domain of science; they belong, he said, to the realm of miracles. That species have a capacity for change is admitted by all evolutionists; but there is no need to invoke modifications other than those represented by ordinary variability. It is well known that in artificial selection this tendency to vary has given rise to numerous distinct races, and there is no reason for denying that it can do the same in nature, by the aid of natural selection. On both lines an advance may be expected with equal probability.

His main argument, however, is that the most striking and most highly adapted modifications may be acquired by successive variations. Each of these may be slight, and they may affect different organs, gradually adapting them to the same purpose. The direction of the adaptations will be determined by the needs in the struggle for life, and natural selection will simply exclude all such changes as occur on opposite or deviating lines. In this way, it is not variability itself which is called upon to explain beautiful adaptations, but it is quite sufficient to suppose that natural selection has operated during long periods in the same way. Eventually, all the acquired characters, being transmitted together, would appear to us, as if they had all been simultaneously developed.

Correlations must play a large part in such special evolutions: when one part is modified, so will be other parts. The distribution of nourishment will come in as one of the causes, the reactions of different organs to the same external influences as another. But no doubt the more effective cause is that of the internal correlations, which, however, are still but dimly understood. Darwin repeatedly laid great stress on this view, although a definite proof of its correctness could not be given in his time. Such proof requires the direct observation of a mutation, and it should be stated here that even the first observations made in this direction have clearly confirmed Darwin’s ideas. The new evening primroses which have sprung in my garden from the old form of Oenothera Lamarckiana, and which have evidently been derived from it, in each case, by a single mutation, do not differ from their parent species in one character only, but in almost all their organs and qualities. Oenothera gigas, for example, has stouter stems and denser foliage; the leaves are larger and broader; its thick flower-buds produce gigantic flowers, but only small fruits with large seeds. Correlative changes of this kind are seen in all my new forms, and they lend support to the view that in the gradual development of highly adapted structures, analogous correlations may have played a large part. They easily explain large deviations from an original type, without requiring the assumption of too many steps.

Monstrosities, as their name implies, are widely different in character from natural species; they cannot, therefore, be adduced as evidence in the investigation of the origin of species. There is no doubt that they may have much in common as regards their manner of origin, and that the origin of species, once understood, may lead to a better understanding of the monstrosities. But the reverse is not true, at least not as regards the main lines of development. Here, it is clear, monstrosities cannot have played a part of any significance.

Reversions, or atavistic changes, would seem to give a better support to the theory of descent through modifications. These have been of paramount importance on many lines of evolution of the animal as well as of the vegetable kingdom. It is often assumed that monocotyledons are descended from some lower group of dicotyledons, probably allied to that which includes the buttercup family. On this view the monocotyledons must be assumed to have lost the cambium and all its influence on secondary growth, the differentiation of the flower into calyx and corolla, the second cotyledon or seed-leaf and several other characters. Losses of characters such as these may have been the result of abrupt changes, but this does not prove that the characters themselves have been produced with equal suddenness. On the contrary, Darwin shows very convincingly that a modification may well be developed by a series of steps, and afterwards suddenly disappear. Many monstrosities, such as those represented by twisted stems, furnish direct proofs in support of this view, since they are produced by the loss of one character and this loss implies secondary changes in a large number of other organs and qualities.

Darwin criticises in detail the hypothesis of great and abrupt changes and comes to the conclusion that it does not give even a shadow of an explanation of the origin of species. It is as improbable as it is unnecessary.

Sports and spontaneous variations must now be considered. It is well known that they have produced a large number of fine horticultural varieties. The cut-leaved maple and many other trees and shrubs with split leaves are known to have been produced at a single step; this is true in the case of the single-leaf strawberry plant and of the laciniate variety of the greater celandine: many white flowers, white or yellow berries and numerous other forms had a similar origin. But changes such as these do not come under the head of adaptations, as they consist for the most part in the loss of some quality or organ belonging to the species from which they were derived. Darwin thinks it impossible to attribute to this cause the innumerable structures, which are so well adapted to the habits of life of each species. At the present time we should say that such adaptations require progressive modifications, which are additions to the stock of qualities already possessed by the ancestors, and cannot, therefore, be explained on the ground of a supposed analogy with sports, which are for the most part of a retrogressive nature.

Excluding all these more or less sudden changes, there remains a long series of gradations of variability, but all of these are not assumed by Darwin to be equally fit for the production of new species. In the first place, he disregards all mere temporary variations, such as size, albinism, etc.; further, he points out that very many species have almost certainly been produced by steps, not greater, and probably not very much smaller, than those separating closely related varieties. For varieties are only small species. Next comes the question of polymorphic species: their occurrence seems to have been a source of much doubt and difficulty in Darwin’s mind, although at present it forms one of the main supports of the prevailing explanation of the origin of new species. Darwin simply states that this kind of variability seems to be of a peculiar nature; since polymorphic species are now in a stable condition their occurrence gives no clue as to the mode of origin of new species. Polymorphic species are the expression of the result of previous variability acting on a large scale; but they now simply consist of more or less numerous elementary species, which, as far as we know, do not at present exhibit a larger degree of variability than any other more uniform species. The vernal whitlow-grass (Draba verna) and the wild pansy are the best known examples; both have spread over almost the whole of Europe and are split up into hundreds of elementary forms. These sub-species show no signs of any extraordinary degree of variability, when cultivated under conditions necessary for the exclusion of inter-crossing. Hooker has shown, in the case of some ferns distributed over still wider areas, that the extinction of some of the intermediate forms in such groups would suffice to justify the elevation of the remaining types to the rank of distinct species. Polymorphic species may now be regarded as the link which unites ordinary variability with the historical production of species. But it does not appear that they had this significance for Darwin; and, in fact, they exhibit no phenomena which could explain the processes by which one species has been derived from another. By thus narrowing the limits of the species-producing variability Darwin was led to regard small deviations as the source from which natural selection derives material upon which to act. But even these are not all of the same type, and Darwin was well aware of the fact.

It should here be pointed out that in order to be selected, a change must first have been produced. This proposition, which now seems self-evident, has, however, been a source of much difference of opinion among Darwin’s followers. The opinion that natural selection produces changes in useful directions has prevailed for a long time. In other words, it was assumed that natural selection, by the simple means of singling out, could induce small and useful changes to increase and to reach any desired degree of deviation from the original type. In my opinion this view was never actually held by Darwin. It is in contradiction with the acknowledged aim of all his work,–the explanation of the origin of species by means of natural forces and phenomena only. Natural selection acts as a sieve; it does not single out the best variations, but it simply destroys the larger number of those which are, from some cause or another, unfit for their present environment. In this way it keeps the strains up to the required standard, and, in special circumstances, may even improve them.

Returning to the variations which afford the material for the sieving- action of natural selection, we may distinguish two main kinds. It is true that the distinction between these was not clear at the time of Darwin, and that he was unable to draw a sharp line between them. Nevertheless, in many cases, he was able to separate them, and he often discussed the question which of the two would be the real source of the differentiation of species. Certain variations constantly occur, especially such as are connected with size, weight, colour, etc. They are usually too small for natural selection to act upon, having hardly any influence in the struggle for life: others are more rare, occurring only from time to time, perhaps once or twice in a century, perhaps even only once in a thousand years. Moreover, these are of another type, not simply affecting size, number or weight, but bringing about something new, which may be useful or not. Whenever the variation is useful natural selection will take hold of it and preserve it; in other cases the variation may either persist or disappear.

In his criticism of miscellaneous objections brought forward against the theory of natural selection after the publication of the first edition of “The Origin of Species”, Darwin stated his view on this point very clearly:–“The doctrine of natural selection or the survival of the fittest, which implies that when variations or individual differences of a beneficial nature happen to arise, these will be preserved.” (“Origin of Species” (6th edition), page 169, 1882.) In this sentence the words “HAPPEN TO ARISE” appear to me of prominent significance. They are evidently due to the same general conception which prevailed in Darwin’s Pangenesis hypothesis. (Cf. de Vries, “Intracellulare Pangenesis”, page 73, Jena, 1889, and “Die Mutationstheorie”, I. page 63. Leipzig, 1901.)

A distinction is indicated between ordinary fluctuations which are always present, and such variations as “happen to arise” from time to time. ((I think it right to point out that the interpretation of this passage from the “Origin” by Professor de Vries is not accepted as correct either by Mr Francis Darwin or by myself. We do not believe that Darwin intended to draw any distinction between TWO TYPES of variation; the words “when variations or individual differences of a beneficial nature happen to arise” are not in our opinion meant to imply a distinction between ordinary fluctuations and variations which “happen to arise,” but we believe that “or” is here used in the sense of ALIAS. With the permission of Professor de Vries, the following extract is quoted from a letter in which he replied to the objection raised to his reading of the passage in question:

“As to your remarks on the passage on page 6, I agree that it is now impossible to see clearly how far Darwin went in his distinction of the different kinds of variability. Distinctions were only dimly guessed at by him. But in our endeavour to arrive at a true conception of his view I think that the chapter on Pangenesis should be our leading guide, and that we should try to interpret the more difficult passages by that chapter. A careful and often repeated study of the Pangenesis hypothesis has convinced me that Darwin, when he wrote that chapter, was well aware that ordinary variability has nothing to do with evolution, but that other kinds of variation were necessary. In some chapters he comes nearer to a clear distinction than in others. To my mind the expression ‘happen to arise’ is the sharpest indication of his inclining in this direction. I am quite convinced that numerous expressions in his book become much clearer when looked at in this way.”

The statement in this passage that “Darwin was well aware that ordinary variability has nothing to do with evolution, but that other kinds of variation were necessary” is contradicted by many passages in the “Origin”. A.C.S.)) The latter afford the material for natural selection to act upon on the broad lines of organic development, but the first do not. Fortuitous variations are the species-producing kind, which the theory requires; continuous fluctuations constitute, in this respect, a useless type.

Of late, the study of variability has returned to the recognition of this distinction. Darwin’s variations, which from time to time happen to arise, are MUTATIONS, the opposite type being commonly designed fluctuations. A large mass of facts, collected during the last few decades, has confirmed this view, which in Darwin’s time could only be expressed with much reserve, and everyone knows that Darwin was always very careful in statements of this kind.

From the same chapter I may here cite the following paragraph: “Thus as I am inclined to believe, morphological differences,…such as the arrangement of the leaves, the divisions of the flower or of the ovarium, the position of the ovules, etc.–first appeared in many cases as fluctuating variations, which sooner or later became constant through the nature of the organism and of the surrounding conditions…but NOT THROUGH NATURAL SELECTION (The italics are mine (H. de V.).); for as these morphological characters do not affect the welfare of the species, any slight deviation in them could not have been governed or accumulated through this latter agency.” (“Origin of Species” (6th edition), page 176.) We thus see that in Darwin’s opinion, all small variations had not the same importance. In favourable circumstances some could become constant, but others could not.

Since the appearance of the first edition of “The Origin of Species” fluctuating variability has been thoroughly studied by Quetelet. He discovered the law, which governs all phenomena of organic life falling under this head. It is a very simple law, and states that individual variations follow the laws of probability. He proved it, in the first place, for the size of the human body, using the measurements published for Belgian recruits; he then extended it to various other measurements of parts of the body, and finally concluded that it must be of universal validity for all organic beings. It must hold true for all characters in man, physical as well as intellectual and moral qualities; it must hold true for the plant kingdom as well as for the animal kingdom; in short, it must include the whole living world.

Quetelet’s law may be most easily studied in those cases where the variability relates to measure, number and weight, and a vast number of facts have since confirmed its exactness and its validity for all kinds of organisms, organs and qualities. But if we examine it more closely, we find that it includes just those minute variations, which, as Darwin repeatedly pointed out, have often no significance for the origin of species. In the phenomena, described by Quetelet’s law nothing “happens to arise”; all is governed by the common law, which states that small deviations from the mean type are frequent, but that larger aberrations are rare, the rarer as they are larger. Any degree of variation will be found to occur, if only the number of individuals studied is large enough: it is even possible to calculate before hand, how many specimens must be compared in order to find a previously fixed degree of deviation.

The variations, which from time to time happen to appear, are evidently not governed by this law. They cannot, as yet, be produced at will: no sowings of thousands or even of millions of plants will induce them, although by such means the chance of their occurring will obviously be increased. But they are known to occur, and to occur suddenly and abruptly. They have been observed especially in horticulture, where they are ranged in the large and ill-defined group called sports. Korschinsky has collected all the evidence which horticultural literature affords on this point. (S. Korschinsky, “Heterogenesis und Evolution”, “Flora”, Vol. LXXXIX. pages 240-363, 1901.) Several cases of the first appearance of a horticultural novelty have been recorded: this has always happened in the same way; it appeared suddenly and unexpectedly without any definite relation to previously existing variability. Dwarf types are one of the commonest and most favourite varieties of flowering plants; they are not originated by a repeated selection of the smallest specimens, but appear at once, without intermediates and without any previous indication. In many instances they are only about half the height of the original type, thus constituting obvious novelties. So it is in other cases described by Korschinsky: these sports or mutations are now recognised to be the main source of varieties of horticultural plants.

As already stated, I do not pretend that the production of horticultural novelties is the prototype of the origin of new species in nature. I assume that they are, as a rule, derived from the parent species by the loss of some organ or quality, whereas the main lines of the evolution of the animal and vegetable kingdom are of course determined by progressive changes. Darwin himself has often pointed out this difference. But the saltatory origin of horticultural novelties is as yet the simplest parallel for natural mutations, since it relates to forms and phenomena, best known to the general student of evolution.

The point which I wish to insist upon is this. The difference between small and ever present fluctuations and rare and more sudden variations was clear to Darwin, although the facts known at his time were too meagre to enable a sharp line to be drawn between these two great classes of variability. Since Darwin’s time evidence, which proves the correctness of his view, has accumulated with increasing rapidity. Fluctuations constitute one type; they are never absent and follow the law of chance, but they do not afford the material from which to build new species. Mutations, on the other hand, only happen to occur from time to time. They do not necessarily produce greater changes than fluctuations, but such as may become, or rather are from their very nature, constant. It is this constancy which is the mark of specific characters, and on this basis every new specific character may be assumed to have arisen by mutation.

Some authors have tried to show that the theory of mutation is opposed to Darwin’s views. But this is erroneous. On the contrary, it is in fullest harmony with the great principle laid down by Darwin. In order to be acted upon by that complex of environmental forces, which Darwin has called natural selection, the changes must obviously first be there. The manner in which they are produced is of secondary importance and has hardly any bearing on the theory of descent with modification. (“Life and Letters” II. 125.)

A critical survey of all the facts of variability of plants in nature as well as under cultivation has led me to the conviction, that Darwin was right in stating that those rare beneficial variations, which from time to time happen to arise,–the now so-called mutations–are the real source of progress in the whole realm of the organic world.


All phenomena of animal and plant life are governed by two sets of causes; one of these is external, the other internal. As a rule the internal causes determine the nature of a phenomenon–what an organism can do and what it cannot do. The external causes, on the other hand, decide when a certain variation will occur, and to what extent its features may be developed.

As a very clear and wholly typical instance I cite the cocks-combs (Celosia). This race is distinguished from allied forms by its faculty of producing the well-known broad and much twisted combs. Every single individual possesses this power, but all individuals do not exhibit it in its most complete form. In some cases this faculty may not be exhibited at the top of the main stem, although developed in lateral branches: in others it begins too late for full development. Much depends upon nourishment and cultivation, but almost always the horticulturist has to single out the best individuals and to reject those which do not come up to the standard.

The internal causes are of a historical nature. The external ones may be defined as nourishment and environment. In some cases nutrition is the main factor, as, for instance, in fluctuating variability, but in natural selection environment usually plays the larger part.

The internal or historical causes are constant during the life-time of a species, using the term species in its most limited sense, as designating the so-called elementary species or the units out of which the ordinary species are built up. These historical causes are simply the specific characters, since in the origin of a species one or more of these must have been changed, thus producing the characters of the new type. These changes must, of course, also be due partly to internal and partly to external causes.

In contrast to these changes of the internal causes, the ordinary variability which is exhibited during the life-time of a species is called fluctuating variability. The name mutations or mutating variability is then given to the changes in the specific characters. It is desirable to consider these two main divisions of variability separately.

In the case of fluctuations the internal causes, as well as the external ones, are often apparent. The specific characters may be designated as the mean about which the observed forms vary. Almost every character may be developed to a greater or a less degree, but the variations of the single characters producing a small deviation from the mean are usually the commonest. The limits of these fluctuations may be called wide or narrow, according to the way we look at them, but in numerous cases the extreme on the favoured side hardly surpasses double the value of that on the other side. The degree of this development, for every individual and for every organ, is dependent mainly on nutrition. Better nourishment or an increased supply of food produces a higher development; only it is not always easy to determine which direction is the fuller and which is the poorer one. The differences among individuals grown from different seeds are described as examples of individual variability, but those which may be observed on the same plant, or on cuttings, bulbs or roots derived from one individual are referred to as cases of partial variability. Partial variability, therefore, determines the differences among the flowers, fruits, leaves or branches of one individual: in the main, it follows the same laws as individual variability, but the position of a branch on a plant also determines its strength, and the part it may take in the nourishment of the whole. Composite flowers and umbels therefore have, as a rule, fewer rays on weak branches than on the strong main ones. The number of carpels in the fruits of poppies becomes very small on the weak lateral branches, which are produced towards the autumn, as well as on crowded, and therefore on weakened individuals. Double flowers follow the same rule, and numerous other instances could easily be adduced.

Mutating variability occurs along three main lines. Either a character may disappear, or, as we now say, become latent; or a latent character may reappear, reproducing thereby a character which was once prominent in more or less remote ancestors. The third and most interesting case is that of the production of quite new characters which never existed in the ancestors. Upon this progressive mutability the main development of the animal and vegetable kingdom evidently depends. In contrast to this, the two other cases are called retrogressive and degressive mutability. In nature retrogressive mutability plays a large part; in agriculture and in horticulture it gives rise to numerous varieties, which have in the past been preserved, either on account of their usefulness or beauty, or simply as fancy-types. In fact the possession of numbers of varieties may be considered as the main character of domesticated animals and cultivated plants.

In the case of retrogressive and degressive mutability the internal cause is at once apparent, for it is this which causes the disappearance or reappearance of some character. With progressive mutations the case is not so simple, since the new character must first be produced and then displayed. These two processes are theoretically different, but they may occur together or after long intervals. The production of the new character I call premutation, and the displaying mutation. Both of course must have their external as well as their internal causes, as I have repeatedly pointed out in my work on the Mutation Theory. (“Die Mutationstheorie”, 2 vols., Leipzig, 1901.)

It is probable that nutrition plays as important a part among the external causes of mutability as it does among those of fluctuating variability. Observations in support of this view, however, are too scanty to allow of a definite judgment. Darwin assumed an accumulative influence of external causes in the case of the production of new varieties or species. The accumulation might be limited to the life-time of a single individual, or embrace that of two or more generations. In the end a degree of instability in the equilibrium of one or more characters might be attained, great enough for a character to give way under a small shock produced by changed conditions of life. The character would then be thrown over from the old state of equilibrium into a new one.

Characters which happen to be in this state of unstable equilibrium are called mutable. They may be either latent or active, being in the former case derived from old active ones or produced as new ones (by the process, designated premutation). They may be inherited in this mutable condition during a long series of generations. I have shown that in the case of the evening primrose of Lamarck this state of mutability must have existed for at least half a century, for this species was introduced from Texas into England about the year 1860, and since then all the strains derived from its first distribution over the several countries of Europe show the same phenomena in producing new forms. The production of the dwarf evening primrose, or Oenothera nanella, is assumed to be due to one of the factors, which determines the tall stature of the parent form, becoming latent; this would, therefore, afford an example of retrogressive mutation. Most of the other types of my new mutants, on the other hand, seem to be due to progressive mutability.

The external causes of this curious period of mutability are as yet wholly unknown and can hardly be guessed at, since the origin of the Oenothera Lamarckiana is veiled in mystery. The seeds, introduced into England about 1860, were said to have come from Texas, but whether from wild or from cultivated plants we do not know. Nor has the species been recorded as having been observed in the wild condition. This, however, is nothing peculiar. The European types of Oenothera biennis and O. muricata are in the same condition. The first is said to have been introduced from Virginia, and the second from Canada, but both probably from plants cultivated in the gardens of these countries. Whether the same elementary species are still growing on those spots is unknown, mainly because the different sub-species of the species mentioned have not been systematically studied and distinguished.

The origin of new species, which is in part the effect of mutability, is, however, due mainly to natural selection. Mutability provides the new characters and new elementary species. Natural selection, on the other hand, decides what is to live and what to die. Mutability seems to be free, and not restricted to previously determined lines. Selection, however, may take place along the same main lines in the course of long geological epochs, thus directing the development of large branches of the animal and vegetable kingdom. In natural selection it is evident that nutrition and environment are the main factors. But it is probable that, while nutrition may be one of the main causes of mutability, environment may play the chief part in the decisions ascribed to natural selection. Relations to neighbouring plants and to injurious or useful animals, have been considered the most important determining factors ever since the time when Darwin pointed out their prevailing influence.

From this discussion of the main causes of variability we may derive the proposition that the study of every phenomenon in the field of heredity, of variability, and of the origin of new species will have to be considered from two standpoints; on one hand we have the internal causes, on the other the external ones. Sometimes the first are more easily detected, in other cases the latter are more accessible to investigation. But the complete elucidation of any phenomenon of life must always combine the study of the influence of internal with that of external causes.


One of the propositions of Darwin’s theory of the struggle for life maintains that the largest amount of life can be supported on any area, by great diversification or divergence in the structure and constitution of its inhabitants. Every meadow and every forest affords a proof of this thesis. The numerical proportion of the different species of the flora is always changing according to external influences. Thus, in a given meadow, some species will flower abundantly in one year and then almost disappear, until, after a series of years, circumstances allow them again to multiply rapidly. Other species, which have taken their places, will then become rare. It follows from this principle, that notwithstanding the constantly changing conditions, a suitable selection from the constituents of a meadow will ensure a continued high production. But, although the principle is quite clear, artificial selection has, as yet, done very little towards reaching a really high standard.

The same holds good for cereals. In ordinary circumstances a field will give a greater yield, if the crop grown consists of a number of sufficiently differing types. Hence it happens that almost all older varieties of wheat are mixtures of more or less diverging forms. In the same variety the numerical composition will vary from year to year, and in oats this may, in bad years, go so far as to destroy more than half of the harvest, the wind-oats (Avena fatua), which scatter their grain to the winds as soon as it ripens, increasing so rapidly that they assume the dominant place. A severe winter, a cold spring and other extreme conditions of life will destroy one form more completely than another, and it is evident that great changes in the numerical composition of the mixture may thus be brought about.

This mixed condition of the common varieties of cereals was well known to Darwin. For him it constituted one of the many types of variability. It is of that peculiar nature to which, in describing other groups, he applies the term polymorphy. It does not imply that the single constituents of the varieties are at present really changing their characters. On the other hand, it does not exclude the possibility of such changes. It simply states that observation shows the existence of different forms; how these have originated is a question which it does not deal with. In his well- known discussion of the variability of cereals, Darwin is mainly concerned with the question, whether under cultivation they have undergone great changes or only small ones. The decision ultimately depends on the question, how many forms have originally been taken into cultivation. Assuming five or six initial species, the variability must be assumed to have been very large, but on the assumption that there were between ten and fifteen types, the necessary range of variability is obviously much smaller. But in regard to this point, we are of course entirely without historical data.

Few of the varieties of wheat show conspicuous differences, although their number is great. If we compare the differentiating characters of the smaller types of cereals with those of ordinary wild species, even within the same genus or family, they are obviously much less marked. All these small characters, however, are strictly inherited, and this fact makes it very probable that the less obvious constituents of the mixtures in ordinary fields must be constant and pure as long as they do not intercross. Natural crossing is in most cereals a phenomenon of rare occurrence, common enough to admit of the production of all possible hybrid combinations, but requiring the lapse of a long series of years to reach its full effect.

Darwin laid great stress on this high amount of variability in the plants of the same variety, and illustrated it by the experience of Colonel Le Couteur (“On the Varieties, Properties, and Classification of Wheat”, Jersey, 1837.) on his farm on the isle of Jersey, who cultivated upwards of 150 varieties of wheat, which he claimed were as pure as those of any other agriculturalist. But Professor La Gasca of Madrid, who visited him, drew attention to aberrant ears, and pointed out, that some of them might be better yielders than the majority of plants in the crop, whilst others might be poor types. Thence he concluded that the isolation of the better ones might be a means of increasing his crops. Le Couteur seems to have considered the constancy of such smaller types after isolation as absolutely probable, since he did not even discuss the possibility of their being variable or of their yielding a changeable or mixed progeny. This curious fact proves that he considered the types, discovered in his fields by La Gasca to be of the same kind as his other varieties, which until that time he had relied upon as being pure and uniform. Thus we see, that for him, the variability of cereals was what we now call polymorphy. He looked through his fields for useful aberrations, and collected twenty-three new types of wheat. He was, moreover, clear about one point, which, on being rediscovered after half a century, has become the starting-point for the new Swedish principle of selecting agricultural plants. It was the principle of single-ear sowing, instead of mixing the grains of all the selected ears together. By sowing each ear on a separate plot he intended not only to multiply them, but also to compare their value. This comparison ultimately led him to the choice of some few valuable sorts, one of which, the “Bellevue de Talavera,” still holds its place among the prominent sorts of wheat cultivated in France. This variety seems to be really a uniform type, a quality very useful under favourable conditions of cultivation, but which seems to have destroyed its capacity for further improvement by selection.

The principle of single-ear sowing, with a view to obtain pure and uniform