ection。
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 guesswithout knowing the factswhat 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