Darwin and Modern Science Part 6

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 a.s.sume it also in regard to ascending variations, as the phenomena of artificial selection certainly justify us in doing. If the j.a.panese breeders were able to lengthen the tail feathers of the c.o.c.k 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 pa.s.sive 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 pa.s.sive flow of the nutritive stream if it is to be beyond the danger of unfavourable variations, or how far it must be weakened pa.s.sively 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 a.s.sumed 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 s.e.xual 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 b.u.t.terflies, 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 p.r.o.nouncedly 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 a.s.sUME 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 a.s.sume 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.

IV. DERIVATIVES OF THE THEORY OF 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.

V. ARGUMENTS FOR THE REALITY OF THE PROCESSES OF SELECTION.

(a) s.e.xUAL SELECTION.

s.e.xual selection goes hand in hand with natural selection. From the very first I have regarded s.e.xual 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 p.r.o.nounced 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 s.e.xual 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 s.e.xual 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 ma.s.s of facts, which, taken together, yield overwhelming evidence of the validity of the principle of s.e.xual 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. s.e.xUAL 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 s.e.x, usually the male, for the possession of the other, usually the female. Its influence can therefore only DIRECTLY affect one s.e.x, in that it equips it better for attaining possession of the other. But the effect may extend indirectly to the female s.e.x, and thus the whole species may be modified, without, however, becoming any more capable of resistance in the struggle for existence, for s.e.xual 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 s.e.x: the manifold "secondary s.e.xual 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 s.e.xual selection is the unequal number of individuals in the two s.e.xes, for if every male immediately finds his mate there can be no compet.i.tion for the possession of the female. Darwin has shown that, for the most part, the inequality between the s.e.xes 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 s.e.xual 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 compet.i.tion for them must take place, and that, when the insects of both s.e.xes 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 pa.s.sed 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 a.s.sume 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 s.e.x. 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 s.e.xual selection, for the union of the s.e.xes 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 s.e.xual 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 s.e.xual 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 s.e.xual excitement in the female became a.s.sociated 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 s.e.xual 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 s.e.x, 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 b.u.t.terflies. Since Fritz Muller found out that certain Brazilian b.u.t.terflies 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 s.e.xes 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 s.e.xes 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 b.u.t.terfly 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 s.e.xual 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 s.e.xual 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 b.u.t.terfly, 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 b.u.t.terflies are "more beautiful" than the females, and in the Tropics in particular they s.h.i.+ne and glow in the most superb colours. I really see no reason why we should doubt the power of s.e.xual selection, and I myself stand wholly on Darwin's side. Even though we certainly cannot a.s.sume 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 b.u.t.terflies. 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 b.u.t.terflies 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 b.u.t.terflies, 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 s.e.xual 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 a.n.a.logy with other secondary s.e.xual characters. If the song of birds and the chirping of the cricket have been evolved through s.e.xual 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 b.u.t.terflies 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 courts.h.i.+p. 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 b.u.t.terflies, 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 s.e.xual 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 courts.h.i.+p-characters, such as coloured spots, decorative feathers, h.o.r.n.y 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 ill.u.s.trated 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 s.e.xual characters, we can hardly fail to be convinced of the truth of the principle of s.e.xual selection. And certainly no one who has accepted natural selection should reject s.e.xual 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.

(b) NATURAL SELECTION.

An actual proof of the theory of s.e.xual 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 a.s.sume that slight intensifications of them give any advantage, and we MUST a.s.sume this SINCE OTHERWISE SECONDARY s.e.xUAL 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 a.s.sociation" (Bristol, 1898), London, 1899, pages 906-909.) were made with 600 pupae of Vanessa urticae, the "tortoise-sh.e.l.l b.u.t.terfly." 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 mult.i.tude of phenomena which cannot be explained in any other way. To this category belong all structures which are only Pa.s.sIVELY 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 a.s.sumption 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-b.u.t.terflies, Kallima inachis. In the single b.u.t.terfly-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 b.u.t.terfly 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 b.u.t.terflies 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-b.u.t.terflies, 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 b.u.t.terfly 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 b.u.t.terfly" (Grapta C.

alb.u.m). 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 s.h.i.+nes through it.

The utility obviously lies in presenting to the bird the very familiar picture of a broken leaf with a clear s.h.i.+ning 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 b.u.t.terflies 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 att.i.tude of the wings in the resting position of the two groups.