The Descent of Man, and Selection in Relation to Sex

As I hear from Sir Andrew Smith, the lion in South Africa sometimes lives with a single female, but generally with more, and, in one case, was found with as many as five females; so that he is polygamous. As far as I can discover, he is the only polygamist amongst all the terrestrial Carnivora, and he alone presents well-marked sexual characters. If, however, we turn to the marine Carnivora, as we shall hereafter see, the case is widely different; for many species of seals offer extraordinary sexual differences, and they are eminently polygamous. Thus, according to Peron, the male sea-elephant of the Southern Ocean always possesses several females, and the sea-lion of Forster is said to be surrounded by from twenty to thirty females. In the North, the male sea-bear of Steller is accompanied by even a greater number of females. It is an interesting fact, as Dr. Gill remarks (15. 'The Eared Seals,' American Naturalist, vol. iv. Jan. 1871.), that in the monogamous species, "or those living in small communities, there is little difference in size between the males and females; in the social species, or rather those of which the males have harems, the males are vastly larger than the females."

Amongst birds, many species, the sexes of which differ greatly from each other, are certainly monogamous. In Great Britain we see well-marked sexual differences, for instance, in the wild-duck which pairs with a single female, the common blackbird, and the bullfinch which is said to pair for life. I am informed by Mr. Wallace that the like is true of the Chatterers or Cotingidae of South America, and of many other birds. In several groups I have not been able to discover whether the species are polygamous or monogamous. Lesson says that birds of paradise, so remarkable for their sexual differences, are polygamous, but Mr. Wallace doubts whether he had sufficient evidence. Mr. Salvin tells me he has been led to believe that humming-birds are polygamous. The male widow-bird, remarkable for his caudal plumes, certainly seems to be a polygamist. (16. 'The Ibis,' vol. iii. 1861, p. 133, on the Progne Widow-bird. See also on the Vidua axillaris, ibid. vol. ii. 1860, p. 211. On the polygamy of the Capercailzie and Great Bustard, see L. Lloyd, 'Game Birds of Sweden,' 1867, pp. 19, and 182. Montagu and Selby speak of the Black Grouse as polygamous and of the Red Grouse as monogamous.) I have been assured by Mr. Jenner Weir and by others, that it is somewhat common for three starlings to frequent the same nest; but whether this is a case of polygamy or polyandry has not been ascertained.

The Gallinaceae exhibit almost as strongly marked sexual differences as birds of paradise or humming-birds, and many of the species are, as is well known, polygamous; others being strictly monogamous. What a contrast is presented between the sexes of the polygamous peacock or pheasant, and the monogamous guinea-fowl or partridge! Many similar cases could be given, as in the grouse tribe, in which the males of the polygamous capercailzie and black-cock differ greatly from the females; whilst the sexes of the monogamous red grouse and ptarmigan differ very little. In the Cursores, except amongst the bustards, few species offer strongly-marked sexual differences, and the great bustard (Otis tarda) is said to be polygamous. With the Grallatores, extremely few species differ sexually, but the ruff (Machetes pugnax) affords a marked exception, and this species is believed by Montagu to be a polygamist. Hence it appears that amongst birds there often exists a close relation between polygamy and the development of strongly-marked sexual differences. I asked Mr. Bartlett, of the Zoological Gardens, who has had very large experience with birds, whether the male tragopan (one of the Gallinaceae) was polygamous, and I was struck by his answering, "I do not know, but should think so from his splendid colours."

It deserves notice that the instinct of pairing with a single female is easily lost under domestication. The wild-duck is strictly monogamous, the domestic-duck highly polygamous. The Rev. W.D. Fox informs me that out of some half-tamed wild-ducks, on a large pond in his neighbourhood, so many mallards were shot by the gamekeeper that only one was left for every seven or eight females; yet unusually large broods were reared. The guinea-fowl is strictly monogamous; but Mr. Fox finds that his birds succeed best when he keeps one cock to two or three hens. Canary-birds pair in a state of nature, but the breeders in England successfully put one male to four or five females. I have noticed these cases, as rendering it probable that wild monogamous species might readily become either temporarily or permanently polygamous.

Too little is known of the habits of reptiles and fishes to enable us to speak of their marriage arrangements. The stickle-back (Gasterosteus), however, is said to be a polygamist (17. Noel Humphreys, 'River Gardens,' 1857.); and the male during the breeding-season differs conspicuously from the female.

To sum up on the means through which, as far as we can judge, sexual selection has led to the development of secondary sexual characters. It has been shewn that the largest number of vigorous offspring will be reared from the pairing of the strongest and best-armed males, victorious in contests over other males, with the most vigorous and best-nourished females, which are the first to breed in the spring. If such females select the more attractive, and at the same time vigorous males, they will rear a larger number of offspring than the retarded females, which must pair with the less vigorous and less attractive males. So it will be if the more vigorous males select the more attractive and at the same time healthy and vigorous females; and this will especially hold good if the male defends the female, and aids in providing food for the young. The advantage thus gained by the more vigorous pairs in rearing a larger number of offspring has apparently sufficed to render sexual selection efficient. But a large numerical preponderance of males over females will be still more efficient; whether the preponderance is only occasional and local, or permanent; whether it occurs at birth, or afterwards from the greater destruction of the females; or whether it indirectly follows from the practice of polygamy.

THE MALE GENERALLY MORE MODIFIED THAN THE FEMALE

Throughout the animal kingdom, when the sexes differ in external appearance, it is, with rare exceptions, the male which has been the more modified; for, generally, the female retains a closer resemblance to the young of her own species, and to other adult members of the same group. The cause of this seems to lie in the males of almost all animals having stronger passions than the females. Hence it is the males that fight together and sedulously display their charms before the females; and the victors transmit their superiority to their male offspring. Why both sexes do not thus acquire the characters of their fathers, will be considered hereafter. That the males of all mammals eagerly pursue the females is notorious to every one. So it is with birds; but many cock birds do not so much pursue the hen, as display their plumage, perform strange antics, and pour forth their song in her presence. The male in the few fish observed seems much more eager than the female; and the same is true of alligators, and apparently of Batrachians. Throughout the enormous class of insects, as Kirby remarks, "the law is that the male shall seek the female." (18. Kirby and Spence, 'Introduction to Entomology,' vol. iii. 1826, p. 342.) Two good authorities, Mr. Blackwall and Mr. C. Spence Bate, tell me that the males of spiders and crustaceans are more active and more erratic in their habits than the females. When the organs of sense or locomotion are present in the one sex of insects and crustaceans and absent in the other, or when, as is frequently the case, they are more highly developed in the one than in the other, it is, as far as I can discover, almost invariably the male which retains such organs, or has them most developed; and this shews that the male is the more active member in the courtship of the sexes. (19. One parasitic Hymenopterous insect (Westwood, 'Modern Class. of Insects,' vol. ii. p. 160) forms an exception to the rule, as the male has rudimentary wings, and never quits the cell in which it is born, whilst the female has well-developed wings. Audouin believes that the females of this species are impregnated by the males which are born in the same cells with them; but it is much more probable that the females visit other cells, so that close inter-breeding is thus avoided. We shall hereafter meet in various classes, with a few exceptional cases, in which the female, instead of the male, is the seeker and wooer.)

The female, on the other hand, with the rarest exceptions, is less eager than the male. As the illustrious Hunter (20. 'Essays and Observations,' edited by Owen, vol. i. 1861, p. 194.) long ago observed, she generally "requires to be courted;" she is coy, and may often be seen endeavouring for a long time to escape from the male. Every observer of the habits of animals will be able to call to mind instances of this kind. It is shewn by various facts, given hereafter, and by the results fairly attributable to sexual selection, that the female, though comparatively passive, generally exerts some choice and accepts one male in preference to others. Or she may accept, as appearances would sometimes lead us to believe, not the male which is the most attractive to her, but the one which is the least distasteful. The exertion of some choice on the part of the female seems a law almost as general as the eagerness of the male.

We are naturally led to enquire why the male, in so many and such distinct classes, has become more eager than the female, so that he searches for her, and plays the more active part in courtship. It would be no advantage and some loss of power if each sex searched for the other; but why should the male almost always be the seeker? The ovules of plants after fertilisation have to be nourished for a time; hence the pollen is necessarily brought to the female organs – being placed on the stigma, by means of insects or the wind, or by the spontaneous movements of the stamens; and in the Algae, etc., by the locomotive power of the antherozooids. With lowly-organised aquatic animals, permanently affixed to the same spot and having their sexes separate, the male element is invariably brought to the female; and of this we can see the reason, for even if the ova were detached before fertilisation, and did not require subsequent nourishment or protection, there would yet be greater difficulty in transporting them than the male element, because, being larger than the latter, they are produced in far smaller numbers. So that many of the lower animals are, in this respect, analogous with plants. (21. Prof. Sachs ('Lehrbuch der Botanik,' 1870, S. 633) in speaking of the male and female reproductive cells, remarks, "verhält sich die eine bei der Vereinigung activ…die andere erscheint bei der Vereinigung passiv.") The males of affixed and aquatic animals having been led to emit their fertilising element in this way, it is natural that any of their descendants, which rose in the scale and became locomotive, should retain the same habit; and they would approach the female as closely as possible, in order not to risk the loss of the fertilising element in a long passage of it through the water. With some few of the lower animals, the females alone are fixed, and the males of these must be the seekers. But it is difficult to understand why the males of species, of which the progenitors were primordially free, should invariably have acquired the habit of approaching the females, instead of being approached by them. But in all cases, in order that the males should seek efficiently, it would be necessary that they should be endowed with strong passions; and the acquirement of such passions would naturally follow from the more eager leaving a larger number of offspring than the less eager.

The great eagerness of the males has thus indirectly led to their much more frequently developing secondary sexual characters than the females. But the development of such characters would be much aided, if the males were more liable to vary than the females – as I concluded they were – after a long study of domesticated animals. Von Nathusius, who has had very wide experience, is strongly of the same opinion. (22. 'Vorträge uber Viehzucht,' 1872, p. 63.) Good evidence also in favour of this conclusion can be produced by a comparison of the two sexes in mankind. During the Novara Expedition (23. 'Reise der Novara: Anthropolog. Theil,' 1867, ss. 216-269. The results were calculated by Dr. Weisbach from measurements made by Drs. K. Scherzer and Schwarz. On the greater variability of the males of domesticated animals, see my 'Variation of Animals and Plants under Domestication,' vol. ii. 1868, p. 75.) a vast number of measurements was made of various parts of the body in different races, and the men were found in almost every case to present a greater range of variation than the women; but I shall have to recur to this subject in a future chapter. Mr. J. Wood (24. 'Proceedings of the Royal Society,' vol. xvi. July 1868, pp. 519 and 524.), who has carefully attended to the variation of the muscles in man, puts in italics the conclusion that "the greatest number of abnormalities in each subject is found in the males." He had previously remarked that "altogether in 102 subjects, the varieties of redundancy were found to be half as many again as in females, contrasting widely with the greater frequency of deficiency in females before described." Professor Macalister likewise remarks (25. 'Proc. Royal Irish Academy,' vol. x. 1868, p. 123.) that variations in the muscles "are probably more common in males than females." Certain muscles which are not normally present in mankind are also more frequently developed in the male than in the female sex, although exceptions to this rule are said to occur. Dr. Burt Wilder (26. 'Massachusetts Medical Society,' vol. ii. No. 3, 1868, p. 9.) has tabulated the cases of 152 individuals with supernumerary digits, of which 86 were males, and 39, or less than half, females, the remaining 27 being of unknown sex. It should not, however, be overlooked that women would more frequently endeavour to conceal a deformity of this kind than men. Again, Dr. L. Meyer asserts that the ears of man are more variable in form than those of a woman. (27. 'Archiv fur Path. Anat. und Phys.' 1871, p. 488.) Lastly the temperature is more variable in man than in woman. (28. The conclusions recently arrived at by Dr. J. Stockton Hough, on the temperature of man, are given in the 'Pop. Sci. Review,' Jan. 1st, 1874, p. 97.)

The cause of the greater general variability in the male sex, than in the female is unknown, except in so far as secondary sexual characters are extraordinarily variable, and are usually confined to the males; and, as we shall presently see, this fact is, to a certain extent, intelligible. Through the action of sexual and natural selection male animals have been rendered in very many instances widely different from their females; but independently of selection the two sexes, from differing constitutionally, tend to vary in a somewhat different manner. The female has to expend much organic matter in the formation of her ova, whereas the male expends much force in fierce contests with his rivals, in wandering about in search of the female, in exerting his voice, pouring out odoriferous secretions, etc.: and this expenditure is generally concentrated within a short period. The great vigour of the male during the season of love seems often to intensify his colours, independently of any marked difference from the female. (29. Prof. Mantegazza is inclined to believe ('Lettera a Carlo Darwin,' 'Archivio per l'Anthropologia,' 1871, p. 306) that the bright colours, common in so many male animals, are due to the presence and retention by them of the spermatic fluid; but this can hardly be the case; for many male birds, for instance young pheasants, become brightly coloured in the autumn of their first year.) In mankind, and even as low down in the organic scale as in the Lepidoptera, the temperature of the body is higher in the male than in the female, accompanied in the case of man by a slower pulse. (30. For mankind, see Dr. J. Stockton Hough, whose conclusions are given in the 'Popular Science Review,' 1874, p. 97. See Girard's observations on the Lepidoptera, as given in the 'Zoological Record,' 1869, p. 347.) On the whole the expenditure of matter and force by the two sexes is probably nearly equal, though effected in very different ways and at different rates.

From the causes just specified the two sexes can hardly fail to differ somewhat in constitution, at least during the breeding-season; and, although they may be subjected to exactly the same conditions, they will tend to vary in a different manner. If such variations are of no service to either sex, they will not be accumulated and increased by sexual or natural selection. Nevertheless, they may become permanent if the exciting cause acts permanently; and in accordance with a frequent form of inheritance they may be transmitted to that sex alone in which they first appeared. In this case the two sexes will come to present permanent, yet unimportant, differences of character. For instance, Mr. Allen shews that with a large number of birds inhabiting the northern and southern United States, the specimens from the south are darker-coloured than those from the north; and this seems to be the direct result of the difference in temperature, light, etc., between the two regions. Now, in some few cases, the two sexes of the same species appear to have been differently affected; in the Agelaeus phoeniceus the males have had their colours greatly intensified in the south; whereas with Cardinalis virginianus it is the females which have been thus affected; with Quiscalus major the females have been rendered extremely variable in tint, whilst the males remain nearly uniform. (31. 'Mammals and Birds of E. Florida,' pp. 234, 280, 295.)

A few exceptional cases occur in various classes of animals, in which the females instead of the males have acquired well pronounced secondary sexual characters, such as brighter colours, greater size, strength, or pugnacity. With birds there has sometimes been a complete transposition of the ordinary characters proper to each sex; the females having become the more eager in courtship, the males remaining comparatively passive, but apparently selecting the more attractive females, as we may infer from the results. Certain hen birds have thus been rendered more highly coloured or otherwise ornamented, as well as more powerful and pugnacious than the cocks; these characters being transmitted to the female offspring alone.

It may be suggested that in some cases a double process of selection has been carried on; that the males have selected the more attractive females, and the latter the more attractive males. This process, however, though it might lead to the modification of both sexes, would not make the one sex different from the other, unless indeed their tastes for the beautiful differed; but this is a supposition too improbable to be worth considering in the case of any animal, excepting man. There are, however, many animals in which the sexes resemble each other, both being furnished with the same ornaments, which analogy would lead us to attribute to the agency of sexual selection. In such cases it may be suggested with more plausibility, that there has been a double or mutual process of sexual selection; the more vigorous and precocious females selecting the more attractive and vigorous males, the latter rejecting all except the more attractive females. But from what we know of the habits of animals, this view is hardly probable, for the male is generally eager to pair with any female. It is more probable that the ornaments common to both sexes were acquired by one sex, generally the male, and then transmitted to the offspring of both sexes. If, indeed, during a lengthened period the males of any species were greatly to exceed the females in number, and then during another lengthened period, but under different conditions, the reverse were to occur, a double, but not simultaneous, process of sexual selection might easily be carried on, by which the two sexes might be rendered widely different.

We shall hereafter see that many animals exist, of which neither sex is brilliantly coloured or provided with special ornaments, and yet the members of both sexes or of one alone have probably acquired simple colours, such as white or black, through sexual selection. The absence of bright tints or other ornaments may be the result of variations of the right kind never having occurred, or of the animals themselves having preferred plain black or white. Obscure tints have often been developed through natural selection for the sake of protection, and the acquirement through sexual selection of conspicuous colours, appears to have been sometimes checked from the danger thus incurred. But in other cases the males during long ages may have struggled together for the possession of the females, and yet no effect will have been produced, unless a larger number of offspring were left by the more successful males to inherit their superiority, than by the less successful: and this, as previously shewn, depends on many complex contingencies.

Sexual selection acts in a less rigorous manner than natural selection. The latter produces its effects by the life or death at all ages of the more or less successful individuals. Death, indeed, not rarely ensues from the conflicts of rival males. But generally the less successful male merely fails to obtain a female, or obtains a retarded and less vigorous female later in the season, or, if polygamous, obtains fewer females; so that they leave fewer, less vigorous, or no offspring. In regard to structures acquired through ordinary or natural selection, there is in most cases, as long as the conditions of life remain the same, a limit to the amount of advantageous modification in relation to certain special purposes; but in regard to structures adapted to make one male victorious over another, either in fighting or in charming the female, there is no definite limit to the amount of advantageous modification; so that as long as the proper variations arise the work of sexual selection will go on. This circumstance may partly account for the frequent and extraordinary amount of variability presented by secondary sexual characters. Nevertheless, natural selection will determine that such characters shall not be acquired by the victorious males, if they would be highly injurious, either by expending too much of their vital powers, or by exposing them to any great danger. The development, however, of certain structures – of the horns, for instance, in certain stags – has been carried to a wonderful extreme; and in some cases to an extreme which, as far as the general conditions of life are concerned, must be slightly injurious to the male. From this fact we learn that the advantages which favoured males derive from conquering other males in battle or courtship, and thus leaving a numerous progeny, are in the long run greater than those derived from rather more perfect adaptation to their conditions of life. We shall further see, and it could never have been anticipated, that the power to charm the female has sometimes been more important than the power to conquer other males in battle.

LAWS OF INHERITANCE

In order to understand how sexual selection has acted on many animals of many classes, and in the course of ages has produced a conspicuous result, it is necessary to bear in mind the laws of inheritance, as far as they are known. Two distinct elements are included under the term "inheritance" – the transmission, and the development of characters; but as these generally go together, the distinction is often overlooked. We see this distinction in those characters which are transmitted through the early years of life, but are developed only at maturity or during old age. We see the same distinction more clearly with secondary sexual characters, for these are transmitted through both sexes, though developed in one alone. That they are present in both sexes, is manifest when two species, having strongly- marked sexual characters, are crossed, for each transmits the characters proper to its own male and female sex to the hybrid offspring of either sex. The same fact is likewise manifest, when characters proper to the male are occasionally developed in the female when she grows old or becomes diseased, as, for instance, when the common hen assumes the flowing tail- feathers, hackles, comb, spurs, voice, and even pugnacity of the cock. Conversely, the same thing is evident, more or less plainly, with castrated males. Again, independently of old age or disease, characters are occasionally transferred from the male to the female, as when, in certain breeds of the fowl, spurs regularly appear in the young and healthy females. But in truth they are simply developed in the female; for in every breed each detail in the structure of the spur is transmitted through the female to her male offspring. Many cases will hereafter be given, where the female exhibits, more or less perfectly, characters proper to the male, in whom they must have been first developed, and then transferred to the female. The converse case of the first development of characters in the female and of transference to the male, is less frequent; it will therefore be well to give one striking instance. With bees the pollen- collecting apparatus is used by the female alone for gathering pollen for the larvae, yet in most of the species it is partially developed in the males to whom it is quite useless, and it is perfectly developed in the males of Bombus or the humble-bee. (32. H. Muller, 'Anwendung der Darwin'schen Lehre,' etc., Verh. d. n. V. Jahrg., xxix. p. 42.) As not a single other Hymenopterous insect, not even the wasp, which is closely allied to the bee, is provided with a pollen-collecting apparatus, we have no grounds for supposing that male bees primordially collected pollen as well as the females; although we have some reason to suspect that male mammals primordially suckled their young as well as the females. Lastly, in all cases of reversion, characters are transmitted through two, three, or many more generations, and are then developed under certain unknown favourable conditions. This important distinction between transmission and development will be best kept in mind by the aid of the hypothesis of pangenesis. According to this hypothesis, every unit or cell of the body throws off gemmules or undeveloped atoms, which are transmitted to the offspring of both sexes, and are multiplied by self-division. They may remain undeveloped during the early years of life or during successive generations; and their development into units or cells, like those from which they were derived, depends on their affinity for, and union with other units or cells previously developed in the due order of growth.