Chapter VI: Darwin’s Theory of Sexual Selection (1)

Sexual Selection

The theory of sexual selection was formulated by Darwin, even in the first edition of the “Origin of Species,” but was developed at much greater length in “The Descent of Man.” “This form of selection depends, not on a struggle for existence in relation to other organic beings or to external conditions, but on a struggle between the individuals of one sex, generally the males, for the possession of the other sex. The result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection. Generally the most vigorous males, those which are best fitted for their place in nature, will leave most progeny. But in many cases victory depends, not so much on general vigor, as on having special weapons, confined to the male sex. A hornless stag or spurless cock would have a poor chance of leaving numerous offspring. Sexual selection, by always allowing the victor to breed, might surely give indomitable courage, length to the spur, and strength to the wing to strike in the spurred leg in nearly the same manner as the brutal cock-fighter by the careful selection of his best cocks.” It is important to notice that the theory of sexual selection is admittedly an extension of the selection principle into a new field. Having accounted for domesticated animals and plants by artificial selection, and for the adaptations of wild species by natural selection, there remained only to account for the secondary sexual differences between the sexes by the principle of sexual selection.

There are two ways in which Darwin supposes sexual selection to act: (1) through competition of the individuals of the same sex with each other,—the strongest or best-equipped for fighting or for finding the individuals of the other sex gaining an advantage; (2) through selection by the individuals of one sex of certain preferred individuals of the other sex.

The first category is natural selection applied to the members of one sex in competition with each other, although the result does not lead to the death of the unsuccessful individual, but excludes it from leaving progeny. In the second category a new element is introduced, namely, the _selective power_ of the individuals of one sex, usually the female. It is this part that adds a distinctly new element to Darwin’s other two theories of selection, and it is this part that we naturally think of as the theory of sexual selection _par excellence_. Darwin makes, however, no sharp distinction between these two sides of his theory, but includes both under the heading of sexual selection.

In order to get the theory itself before us in as concrete form as possible, let us examine some of the cases that Darwin has given to show how he supposes the process to be carried out.

“There are many other structures and instincts which must have been developed through sexual selection—such as the weapons of offence and the means of defence of the males for fighting with and driving away their rivals—their courage and pugnacity—their various ornaments—their contrivances for producing vocal or instrumental music—and their glands for emitting odors, most of these latter structures serving only to allure or excite the female. It is clear that these characters are the result of sexual and not of ordinary selection, since unarmed, unornamented, or unattractive males would succeed equally well in the battle for life and in leaving a numerous progeny, but for the presence of better-endowed males. We may infer that this would be the case, because the females, which are unarmed and unornamented, are able to survive and procreate their kind. Secondary sexual characters of the kind just referred to will be fully discussed in the following chapters, as being in many respects interesting, but especially as depending on the will, choice, and rivalry of the individuals of either sex. When we behold two males fighting for the possession of the female, or several male birds displaying their gorgeous plumage, and performing strange antics before an assembled body of females, we cannot doubt that, though led by instinct, they know what they are about, and consciously exert their mental and bodily powers.”

This general statement gives an idea of the class of phenomena that Darwin proposes to explain by the theory of sexual selection. The close resemblance between this process and that of artificial selection may be gathered from the following statement:—

“Just as man can improve the breed of his game-cocks by the selection of those birds which are victorious in the cockpit, so it appears that the strongest and most vigorous males, or those provided with the best weapons, have prevailed under nature, and have led to the improvement of the natural breed or species. A slight degree of variability leading to some advantage, however slight, in reiterated deadly contests would suffice for the work of sexual selection; and it is certain that secondary sexual characters are eminently variable. Just as man can give beauty, according to his standard of taste, to his male poultry, or more strictly can modify the beauty originally acquired by the parent species, can give to the Sebright bantam a new and elegant plumage, an erect and peculiar carriage—so it appears that female birds in a state of nature have, by a long selection of the more attractive males, added to their beauty or other attractive qualities. No doubt this implies powers of discrimination and taste on the part of the female which will at first appear extremely improbable; but by the facts to be adduced hereafter, I hope to be able to show that the females actually have these powers. When, however, it is said that the lower animals have a sense of beauty, it must not be supposed that such sense is comparable with that of a cultivated man, with his multiform and complex associated ideas. A more just comparison would be between the taste for the beautiful in animals, and that in the lowest savages, who admire and deck themselves with any brilliant, glittering, or curious object.”

Darwin did not close his eyes to the difficulties which the theory had to contend against. One of the most formidable of these objections is described in the following words: “Our difficulty in regard to sexual selection lies in understanding how it is that the males which conquer other males, or those which prove the most attractive to the females, leave a greater number of offspring to inherit their superiority than their beaten and less attractive rivals. Unless this result does follow, the characters which give to certain males an advantage over others could not be perfected and augmented through sexual selection. When the sexes exist in exactly equal numbers, the worst-endowed males will (except where polygamy prevails) ultimately find females, and leave as many offspring, as well fitted for their general habits of life, as the best-endowed males. From various facts and considerations, I formerly inferred that with most animals, in which secondary sexual characters are well developed, the males considerably exceeded the females in number; but this is not by any means always true. If the males were to the females as two to one, or as three to two, or even in a somewhat lower ratio, the whole affair would be simple; for the better-armed or more attractive males would leave the largest number of offspring. But after investigating, as far as possible, the numerical proportion of the sexes, I do not believe that any great inequality in number commonly exists. In most cases sexual selection appears to have been effective in the following manner.

“Let us take any species, a bird for instance, and divide the females inhabiting a district into two equal bodies, the one consisting of the more vigorous and better-nourished individuals, and the other of the less vigorous and healthy. The former, there can be little doubt, would be ready to breed in the spring before the others; and this is the opinion of Mr. Jenner Weir, who has carefully attended to the habits of birds during many years. There can also be no doubt that the most vigorous, best-nourished and earliest breeders would on an average succeed in rearing the largest number of fine offspring. The males, as we have seen, are generally ready to breed before the females; the strongest, and with some species the best-armed of the males, drive away the weaker; and the former would then unite with the more vigorous and better-nourished females, because they are the first to breed. Such vigorous pairs would surely rear a larger number of offspring than the retarded females, which would be compelled to unite with the conquered and less powerful males, supposing the sexes to be numerically equal; and this is all that is wanted to add, in the course of successive generations, to the size, strength and courage of the males, or to improve their weapons.”

I shall comment later on the points here raised, but we should not let this opportunity pass without noticing, that even if the pairing were to follow according to the method here imagined, still the argument breaks down at the critical point, for there is no evidence that the more precocious females would rear a larger number of offspring than the more normal females, or even those that breed somewhat later.

The greater eagerness of the males which has been observed in so many different classes of animals is accounted for as follows:—

“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.”

Thus we are led to the rather complex conclusion, that the more eager males will leave more descendants, and those that are better endowed with ornaments will be the ones selected. But unless it can be shown that there is some connection between greater eagerness and better ornamentation, it might often occur that the less ornamented were the more eager individuals, in which case there would be an apparent conflict between the two acquirements.

After giving some cases of the greater variability of the males, in respect to characters that are not connected with sexual selection, and presumably not the result of any kind of selection, Darwin concludes: “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 vigor of the male during the season of love seems often to intensify his colors, independently of any marked difference from the female. 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. 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.”

Again: “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 shows that with a large number of birds inhabiting the northern and southern United States, the specimens from the south are darker-colored 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 _Agelæus phœniceus_ the males have had their colors 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.”

The admissions contained in this statement would seem to jeopardize the entire question, for, if it is admitted that, on account of the difference in the constitution of the two sexes, the influence of the surrounding conditions would produce a different effect on them, it would seem that there is no need whatsoever for the theory of sexual selection. What Darwin is probably attempting to show is that the material for the further action of sexual selection is already given; but the question may well be asked, if the external conditions have done so much, why may they not have gone farther and produced the entire result?

Darwin makes the following suggestion to account for those cases in which the female is the more highly colored:—

“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 colors, 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 colored or otherwise ornamented, as well as more powerful and pugnacious than the cocks; these characters being transmitted to the female offspring alone.”

Then follows immediately the discussion as to whether a double process of sexual selection may not be supposed to go on at the same time. “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.”

The improbability of such a process is so manifest that the suggestion can scarcely be looked upon as anything more than pure speculation. We shall have occasion later to return to the same subject, and point out its bearing more explicitly.

Nearly the whole animal kingdom is passed in review by Darwin from the point of view of the sexual selection theory. There is brought together a large number of extremely interesting facts, and if the theory did no more than merely hold them together, it has served, in this respect, a useful end. We may select some of the most instructive cases by way of illustrating the theory.

In many of the lower animals in which the sexes are separated, and these alone, of course, can be supposed to come within the range of the theory, there are no striking differences between the sexes, in regard to ornamentation, although in other respects differences may exist.

“Moreover it is almost certain that these animals have too imperfect senses and much too low mental powers, to appreciate each other’s beauty or other attractions, or to feel rivalry.

“Hence in these classes or subkingdoms, such as the Protozoa, Cœlenterata, Echinodermata, Scolecida, secondary sexual characters, of the kind which we have to consider, do not occur; and this fact agrees with the belief that such characters in the higher classes have been acquired through sexual selection, which depends on the will, desire, and choice of either sex.”

There are some cases, however, in which animals low in the scale show a difference in the ornamentation of the two sexes. A few cases have been recorded in the roundworms, where different shades of the same tint distinguish the sexes. In the annelids the sexes are sometimes so different, that, as Darwin remarks, they have been placed in different genera and even families, “yet the differences do not seem to be of the kind which can be safely attributed to sexual selection.” In regard to the nemertian worms, although they “vie in variety and beauty of coloring with any other group in the invertebrate series,” yet McIntosh states that he “cannot discover that these colors are of any service.” In the copepods, belonging to the group of lower Crustacea, Darwin excludes those cases in which the males alone “are furnished with perfect swimming legs, antennæ, and sense-organs; the females being destitute of these organs, with their bodies often consisting of a mere distorted mass,” because these extraordinary differences between the two sexes are no doubt related to their widely different habits of life. Nevertheless, it is important to observe that such extreme differences may exist in cases where sexual selection cannot come in, because of the absence of eyes in the female.

In regard to another copepod, Saphirina, Darwin points out that the males are furnished with minute scales, which exhibit beautiful changing colors, and these are absent in the females; yet he states that it would be extremely rash to conclude that these curious organs serve to attract the females. Differences in the sexes are also found in one species of Squilla, and a species of Gelasimus. In the latter case Darwin thinks that the difference is probably due to sexual selection. In addition to these cases, recorded by Darwin, there may be added the two remarkable cases, shown in our Figure 2 A, B, of _Calocalanus pavo_, the female of which has a gorgeous tail worthy of a peacock, and of _Calocalanus plumulosus_, in which one of the setæ of the tail is drawn out into a long featherlike structure. In the former, the male is much more modestly adorned, as shown in Figure 2 C; in the latter species the male is unknown.

Fig. 2.—A male of the copepod, _Calocalanus plumulosus_.
B and C, a male and a female of _Calocalanus pavo_. (After
Giesbrecht.)]

In spiders, where as a rule the sexes do not differ much from each other in color, the males are often of a darker shade than the females. “In some species, however, the difference is conspicuous; thus the female of _Sparassus smaragdulus_ is dullish green, whilst the adult male has the abdomen of a fine yellow with three longitudinal stripes of rich red.” Darwin believes that sexual selection must take place in this group, because Canestrini has observed that the males fight for the possession of the females. He has also stated that the males pay court to the female, and that she rejects some of the males who court her, and sometimes devours them, until finally one is chosen. Darwin believed, on this evidence, that the difference in color of the sexes had been acquired by sexual selection, “though we have here not the best kind of evidence—the display by the male of his ornaments.” This evidence has, however, now been supplied through the interesting observations of Mr. and Mrs. Peckham. These accurate observers have studied the courtship of the male, and observed that during the process, he twists and turns his body in such a way as to show to best advantage his colors to the female. From their account this certainly appears to be the result of his movements, but whether this is really the case, and whether the female makes any choice amongst her suitors, according to whether they are more or less brilliantly marked, we are absolutely ignorant. The following account given by Darwin should not pass unnoticed:—

“The male is generally much smaller than the female, sometimes to an extraordinary degree, and he is forced to be extremely cautious in making his advances, as the female often carries her coyness to a dangerous pitch. De Geer saw a male that ‘in the midst of his preparatory caresses was seized by the object of his attentions, enveloped by her in a web and then devoured, a sight which, as he adds, filled him with horror and indignation.’ The Rev. O. P. Cambridge accounts in the following manner for the extreme smallness of the male in the genus Nephila. ‘M. Vinson gives a graphic account of the agile way in which the diminutive male escapes from the ferocity of the female, by gliding about and playing hide and seek over her body and along her gigantic limbs: in such a pursuit it is evident that the chances of escape would be in favor of the smallest males, while the larger ones would fall early victims; thus gradually a diminutive race of males would be selected, until at last they would dwindle to the smallest possible size compatible with the exercise of their generative functions,—in fact probably to the size we now see them, _i.e._ so small as to be a sort of parasite upon the female, and either beneath her notice, or too agile and too small for her to catch without great difficulty.’”

It is certainly surprising to find Darwin ascribing even this difference in size between the sexes to the action of selection. Is it not a little ludicrous to suppose that the females have reduced the males to a size too small for them to catch?

There are many cases known in the animal kingdom where there is a difference in size between the two sexes, especially in the group of insects; but I doubt very much if they are to be accounted for as the result of sexual selection. In some of these cases Darwin accounts for the larger size of the female, on account of the large number of eggs which she has to carry. In other insects where the male is larger, as in the stag-beetle, the size is ascribed to the conflicts of the males, leading to the survival of the larger individuals. In still other cases, where the males are larger, but do not fight, an explanation is admittedly wanting; but it is suggested that here there would be no necessity for the males to be smaller than the females in order to mature before them (as is supposed to happen in other species), for in these cases the individuals are not short-lived, and there would be ample time for pairing. Again, although the males of nearly all bees are smaller than the females, yet the reverse is true in those forms in which the females are fertilized during the marriage flight. The explanation offered is that in these forms the male carries the female, and this is assumed to require greater size on his part. This loose way of guessing, as to a possible explanation, is characteristic of the whole hypothesis of sexual selection. First one, and then another, guess is made as to the causes of the differences between the sexes. It is not shown in a single one of the instances that the postulated cause has really had anything to do with the differences in question; and the attempt to show that the theory is probable, by pointing out the large number of cases which it appears to account for, is weakened to a very great degree by the number of exceptional cases, for which an equally ready explanation of a different kind is forthcoming. This way of giving loose rein to the imagination has been the bane of the method that has followed hard on the track of Darwin’s hypothesis, and for which his example has been in no small measure responsible. Thus, in the case just quoted, there are no less than four distinct conjectures made to account for the differences in size between the sexes, and each guess involves an entirely different set of processes. Considering the complicated relation of the life of organisms, it may be doubted if any of the imagined processes could bring about the result, and certainly not a single one has been shown to be a real, or a sufficient, cause in the evolutionary process. Neither the actuality of the postulated causes, nor their application to a particular case, has been shown to exist.

In the Diptera, or flies, Wallace records one interesting case of sexual difference in the genus Elaphomyia of New Guinea, in which the males are furnished with horns, which the females lack. Darwin writes:—

“The horns spring from beneath the eyes, and curiously resemble those of a stag, being either branched or palmated. In one of the species, they equal the whole body in length. They might be thought to be adapted for fighting, but as in one species they are of a beautiful pink color, edged with black, with a pale central stripe, and as these insects have altogether a very elegant appearance, it is perhaps more probable that they serve as ornaments.”

Presumably, therefore, Darwin means these colored horns have been acquired by sexual selection.

In the Hemiptera, or bugs, both sexes of some species are “beautifully colored,” and as the members of the group are often unpalatable to other animals, the color in this case is supposed to act as a warning signal.

In other cases it is stated, however, that “a small pink and green species” could hardly be distinguished from the buds on the trunks of the lime trees which this insect frequents. In this case the color appears “to be directly protective.” Thus without any means of forming a correct judgment, the color of one animal is supposed to be the result of natural selection, since it resembles its surroundings, but of sexual selection if the color is present or more pronounced in one sex. Where neither view can easily be applied, the color is ascribed in a general way to the nature of the organism.

In respect to the group of Hymenoptera, or bees, Darwin records the following cases:—

“In this order slight differences in color, according to sex, are common, but conspicuous differences are rare except in the family of bees; yet both sexes of certain groups are so brilliantly colored—for instance in Chrysis, in which vermilion and metallic greens prevail—that we are tempted to attribute the result to sexual selection. In the Ichneumonidæ, according to Mr. Walsh, the males are almost universally lighter-colored than the females. On the other hand, in the Tenthredinidæ the males are generally darker than the females. In the Siricidæ the sexes frequently differ; thus the male of _Sirex juvencus_ is banded with orange, whilst the female is dark purple; but it is difficult to say which sex is the more ornamented.”

In other families of bees, differences in the color of the sexes have been recorded, and since the males have been seen fighting for the possession(?) of the females, and since bees are known to recognize differences in color, Darwin believes that:—

“In some species the more beautiful males appear to have been selected by the females; and in others the more beautiful females by the males. Consequently in certain genera, the males of the several species differ much in appearance, whilst the females are almost indistinguishable; in other genera the reverse occurs. H. Müller believes that the colors gained by one sex through sexual selection have often been transferred in a variable degree to the other sex, just as the pollen-collecting apparatus of the female has often been transferred to the male, to whom it is absolutely useless.”

Although in beetles the sexes are generally colored alike, yet in some of the longicorns there are exceptions to the rule. “Most of these insects are large and splendidly colored. The males in the genus Pyrodes, which I saw in Mr. Bates’s collection, are generally redder but rather duller than the females, the latter being colored of a more or less splendid golden-green. On the other hand, in one species the male is golden-green, the female being richly tinted with red and purple. In the genus Esmeralda the sexes differ so greatly in color that they have been ranked as distinct species; in one species both are of a beautiful shining green, but the male has a red thorax. On the whole, as far as I could judge, the females of those Prionidæ, in which the sexes differ, are colored more richly than the males, and this does not accord with the common rule in regard to color, when acquired through sexual selection.”

The great horns that rise from the heads of many male beetles are very striking cases of sexual difference, and Darwin compares them to the horns of stags and of the rhinoceros. They “are wonderful from their size and shapes.” Darwin offers the following conjecture as to their meaning: “The extraordinary size of the horns, and their widely different structure in closely allied forms, indicate that they have been formed for some purpose; but their excessive variability in the males of the same species leads to the inference that this purpose cannot be of a definite nature. The horns do not show marks of friction, as if used for any ordinary work. Some authors suppose that as the males wander about much more than the females, they require horns as a defence against their enemies; but as the horns are often blunt, they do not seem well adapted for defence. The most obvious conjecture is that they are used by the males for fighting together; but the males have never been observed to fight; nor could Mr. Bates, after a careful examination of numerous species, find any sufficient evidence, in their mutilated or broken condition, of their having been thus used. If the males had been habitual fighters, the size of their bodies would probably have been increased through sexual selection, so as to have exceeded that of the females; but Mr. Bates, after comparing the two sexes in above a hundred species of the Copridæ, did not find any marked difference in this respect amongst well-developed individuals. In Lethrus, moreover, a beetle belonging to the same great division of the lamellicorns, the males are known to fight, but are not provided with horns, though their mandibles are much larger than those of the female.”

“The conclusion that the horns have been acquired as ornaments is that which best agrees with the fact of their having been so immensely, yet not fixedly, developed,—as shown by their extreme variability in the same species, and by their extreme diversity in closely allied species. This view will at first appear extremely improbable; but we shall hereafter find with many animals standing much higher in the scale, namely fishes, amphibians, reptiles and birds, that various kinds of crests, knobs, horns and combs have been developed apparently for this sole purpose.”

It is asking a great deal to suppose that animals, so dull and sluggish as these beetles, are endowed with a sufficient æsthetic discrimination to select in each generation those males whose horns are a little longer than the average. The resemblance of the horns to those of stags is, as Darwin points out, obvious, but in the latter case also it remains to be proven that they are the result of sexual selection, as Darwin believes to be the case; but the evidence for this belief is not much better, as we shall see in the case of the antlers of deer, than it is in these beetles.

In regard to butterflies, the males and females are both often equally brilliantly colored; in other species the differences in the sexes are very striking. Darwin states:—

“Even within the same genus we often find species presenting extraordinary differences between the sexes, whilst others have their sexes closely alike.” The fine colors of the wings of many moths are also supposed by Darwin to have arisen through sexual selection, although the colors are usually on the lower wings, which are covered during the day by the less ornamented upper wings. It is assumed that, since the moths often begin to fly at dusk, their colors might at this time be seen and appreciated by the other sex. It should not be overlooked, however, that, in the case of some of the most highly colored moths, it is known that the males find the females through the sense of smell. Moreover, although moths are often finely colored, Darwin points out that “it is a singular fact that no British moths which are brilliantly colored, and, as far as I can discover, hardly any foreign species, differ much in color according to sex; though this is the case with many brilliant butterflies.”

Yet Darwin does not hesitate to conclude: “From the several foregoing facts it is impossible to admit that the brilliant colors of butterflies, and of some few moths, have commonly been acquired for the sake of protection. We have seen that their colors and elegant patterns are arranged and exhibited as if for display. Hence I am led to believe that the females prefer or are most excited by the more brilliant males; for on any other supposition the males would, as far as we can see, be ornamented to no purpose. We know that ants and certain lamellicorn beetles are capable of feeling an attachment for each other, and that ants recognize their fellows after an interval of several months. Hence there is no abstract improbability in the Lepidoptera, which probably stand nearly or quite as high in the scale as these insects, having sufficient mental capacity to admire bright colors. They certainly discover flowers by color.”

So far as the evidence of ants having an attachment for each other is concerned, we may eliminate this part of the argument, since the evidence on which the statement is based is now regarded as only showing that ants recognize each other by their sense of smell, which resides in the antennæ. Hence the so-called fondling means only that the ants are trying by smell to determine the odor of the other individual.

Darwin points out a number of cases in which the females are more brightly colored than the males, and for such cases he reverses the process of selection, supposing that the males have been discriminating, and have not “gladly accepted any female.” No explanation is offered to account for this reversal of instinct, in fact, no evidence to show that such a reversal really exists. Darwin points out that in most cases the male insect carries the female during the period of union, while in two species of butterflies, _Colias edusa_ and _hyale_, the females carry the males, and the females are here the more highly colored. He suggests that since in this case “the females take the more active part in the final marriage ceremony, so we may suppose that they likewise do so in the wooing; and in this case we can understand how it is that they have been rendered the more beautiful.”

A most significant fact in connection with the difference in sexual coloration of butterflies did not escape Darwin’s attention.

“Whilst reflecting on the beauty of many butterflies, it occurred to me that some caterpillars were splendidly colored; and as sexual selection could not possibly have here acted, it appeared rash to attribute the beauty of the mature insect to this agency, unless the bright colors of their larvæ could be somehow explained. In the first place, it may be observed that the colors of caterpillars do not stand in any close correlation with those of the mature insect. Secondly, their bright colors do not serve in any ordinary manner as a protection. Mr. Bates informs me, as an instance of this, that the most conspicuous caterpillar which he ever beheld (that of a Sphinx) lived on the large green leaves of a tree on the open llanos of South America; it was about four inches in length, transversely banded with black and yellow, and with its head, legs, and tail of a bright red. Hence it caught the eye of any one who passed by, even at the distance of many yards, and no doubt that of every passing bird.”

Darwin applied to Wallace for a solution of this difficulty, and received the reply that he “thought it probable that conspicuously colored caterpillars were protected by having a nauseous taste; but as their skin is extremely tender, and as their intestines readily protrude from a wound, a slight peck from the beak of a bird would be as fatal to them as if they had been devoured. Hence, as Mr. Wallace remarks, ‘distastefulness alone would be insufficient to protect a caterpillar unless some outward sign indicated to its would-be destroyer that its prey was a disgusting morsel.’ Under these circumstances it would be highly advantageous to a caterpillar to be instantaneously and certainly recognized as unpalatable by all birds and other animals. Thus the most gaudy colors would be serviceable, and might have been gained by variation and the survival of the most easily recognized individuals.”

It need scarcely be pointed out that an occasional peck can scarcely be supposed to have led to the splendid development of color shown by some caterpillars, and Darwin confesses that at first sight this hypothesis appears bold, but nevertheless he believes that it will be found to be true. He adds, “We cannot, however, at present thus explain the elegant diversity in the colors of many caterpillars.”

A most important fact in this connection should not be overlooked, namely, that in the caterpillar stage the sexual organs are so little developed that it is generally impossible at this time to distinguish between the sexes, unless a microscopic examination is made. This gives us, perhaps, a clew as to the difference between the mature sexual forms. These differences are connected with difference of sex itself. This conclusion also fits in well with the fact that during the period when the sexual organs are at the height of their development the individuals are most brilliantly colored. The primary cause of the brilliant color of many animals concerns us here only secondarily, for, since it is known that many of the lower forms are no less brilliantly and elaborately colored than are the sexes of the higher forms, it is not surprising that the sexes themselves sometimes differ in this respect.

Organs for producing sounds of different sorts are present in some insects, and these organs Darwin includes under the head of secondary sexual organs. In the group of Hemiptera, or bugs, the cicadas are the most familiar species that produce sounds. The noise is made by the males; the females are quite mute.

“With respect to the object of the music, Dr. Hartman, in speaking of the _Cicada septemdecim_ of the United States, says, ‘the drums are now (June 6th and 7th, 1851) heard in all directions. This I believe to be the marital summons from the males. Standing in thick chestnut sprouts about as high as my head, where hundreds were around me, I observed the females coming around the drumming males.’ He adds, ‘this season (August, 1868) a dwarf pear-tree in my garden produced about fifty larvæ of _C. pruinosa_; and I several times noticed the females to alight near a male while he was uttering his clanging notes.’ Fritz Müller writes to me from S. Brazil that he has often listened to a musical contest between two or three males of a species with a particularly loud voice, seated at a considerable distance from each other: as soon as one had finished his song, another immediately began, and then another. As there is so much rivalry between the males, it is probable that the females not only find them by their sounds, but that, like female birds, they are excited or allured by the male with the most attractive voice.”

In the flies the following cases are given by Darwin:—

“That the males of some Diptera fight together is certain; for Professor Westwood has several times seen this with the Tipulæ. The males of other Diptera apparently try to win the females by their music: H. Müller watched for some time two males of an Eristalis courting a female; they hovered above her, and flew from side to side, making a high humming noise at the same time. Gnats and mosquitoes (Culicidæ) also seem to attract each other by humming; and Professor Mayer has recently ascertained that the hairs on the antennæ of the male vibrate in unison with the notes of a tuning-fork, within the range of the sounds emitted by the female.”

In the crickets, grasshoppers, and locusts, the males “are remarkable for their musical powers”; and it is generally assumed that the sounds serve to call or to excite the female. In these forms the noise is made by rubbing the wings over each other or the legs against the wing-covers.

In some of these forms both sexes have stridulating organs, and in one case they differ to a certain extent from each other. “Hence we cannot suppose that they have been transferred from the male to the female, as appears to have been the case with the secondary sexual characters of many other animals. They must have been independently developed in the two sexes, which no doubt mutually call to each other during the season of love.”

Some beetles also possess rasping organs in different parts of the body, but they cannot produce much noise by this means.

“We thus see that in the different coleopterous families the stridulating organs are wonderfully diversified in position, but not much in structure. Within the same family some species are provided with these organs, and others are destitute of them. This diversity is intelligible, if we suppose that originally various beetles made a shuffling or hissing noise by the rubbing together of any hard and rough parts of their bodies, which happened to be in contact; and that from the noise thus produced being in some way useful, the rough surfaces were gradually developed into regular stridulating organs. Some beetles as they move, now produce, either intentionally or unintentionally, a shuffling noise, without possessing any proper organs for the purpose.”

Darwin says that he expected from analogy to find in this group also differences in the sexes, but none such were found, although in some cases the males alone possess certain characters or have them more highly developed.

It is important not to forget, when considering all questions connected with sexual selection, that in order for the result to be successful it is not only necessary that the female respond to the noises and music of the other sex, but that she choose the suitor that makes the greatest, or the most pleasing, noise. If the stridulating organs are only used by the animals in finding each other, then the case might be considered as coming under the head of natural selection. If this be granted, then it may be claimed, and apparently Darwin is inclined to adopt this view, that those males that make the most noise will be more likely to be heard, and possibly approached. They will, therefore, be more likely to leave descendants. We have already considered this question when dealing with the theory of natural selection in the preceding chapter and need not go over the ground again. This much may, however, be said again, that even if it is probable that these organs are of use to the animals in finding each other, and this seems not improbable, it does not follow that the organs have been acquired through selection for this purpose.

Darwin finds his best examples of secondary sexual characters in the group of vertebrates, and since in this group the intelligence is of a higher order than in the other groups, the argument that the female chooses the more pleasing suitor is made to appear more plausible.

The elongation of the lower jaw that occurs in a few fishes at the breeding season is regarded as a secondary sexual character. On the other hand, Darwin recognizes the following difficulty in regard to the size of the males:—

“In regard to size, M. Carbonnier maintains that the female of almost all fishes is larger than the male; and Dr. Günther does not know of a single instance in which the male is actually larger than the female. With some cyprinodonts the male is not even half as large. As in many kinds of fishes the males habitually fight together, it is surprising that they have not generally become larger and stronger than the females through the effects of sexual selection. The males suffer from their small size, for, according to M. Carbonnier, they are liable to be devoured by the females of their own species when carnivorous, and no doubt by other species. Increased size must be in some manner of more importance to the females, than strength and size are to the males for fighting with other males; and this perhaps is to allow of the production of a vast number of ova.”

The last sentence implies that this particular case is to be explained by the females becoming larger on account of the number of eggs that they are to produce. But why was not the same explanation offered in the case of the spiders? It is this uncertain way of applying any explanation that suggests itself, that puts the whole method in an unfortunate light.

In many species of fish the males are brighter in color than the females. In the case of _Callionymus lyra_, Darwin states:—

“When fresh caught from the sea the body is yellow of various shades, striped and spotted with vivid blue on the head; the dorsal fins are pale brown with dark longitudinal bands, the ventral, caudal, and anal fins being bluish black. The female, or sordid dragonet, was considered by Linnæus, and by many subsequent naturalists, as a distinct species; it is of a dingy reddish brown, with the dorsal fin brown and the other fins white. The sexes differ also in the proportional size of the head and mouth, and in the position of the eyes; but the most striking difference is the extraordinary elongation in the male of the dorsal fin. Mr. W. Saville Kent remarks that this ‘singular appendage appears from my observations of the species in confinement, to be subservient to the same end as the wattles, crests, and other abnormal adjuncts of the male in gallinaceous birds, for the purpose of fascinating their mates.’”

In the case of another fish, _Cottus scorpius_, there is also a great difference between the sexes, and here the males become very brilliant only at the breeding season. In other fishes, in which the sexes are colored alike, the males may become more brilliant during the breeding season. This, too, is explained by Darwin on the assumption that those males that have varied at the breeding season, so as to become more brightly colored, have been chosen in preference to the other males.

A few cases are cited in which it has been observed that the males appear to exhibit themselves before the females, as in the following case of the Chinese Macropus:—

“The males are most beautifully colored, more so than the females. During the breeding season they contend for the possession of the females; and, in the act of courtship, expand their fins, which are spotted and ornamented with brightly colored rays, in the same manner, according to M. Carbonnier, as the peacock. They then also bound about the females with much vivacity, and appear by ‘l’étalage de leurs vives couleurs chercher à attirer l’attention des femelles, lesquelles ne paraissaient indifférentes à ce manége, elles nageaient avec une molle lenteur vers les mâles et semblaient se complaire dans leur voisinage.’”

In this connection Darwin makes the following general statement:—

“The males sedulously court the females, and in one case, as we have seen, take pains in displaying their beauty before them. Can it be believed that they would thus act to no purpose during their courtship? And this would be the case, unless the females exert some choice and select those males which please or excite them most. If the female exerts such choice, all the above facts on the ornamentation of the males become at once intelligible by the aid of sexual selection.”

While it may readily be granted that display of the male may have for its purpose the excitement of the female, it is another question as to whether she will be more excited by the more beautiful suitor. The attentions of the male may be supposed to have a purpose, even if the female does not choose the more beautiful of her suitors. It is this last proposition, so necessary for the theory of sexual selection, that seems improbable. But even if it were probable, there are, as we shall see, other difficulties to be overcome before we should be justified in accepting Darwin’s statement quoted above, concerning the results of sexual selection.

In regard to those species of fish in which both sexes are equally ornamented, Darwin returns once more to his hypothesis that the color of the male, acquired through sexual selection, may be transmitted to the other sex, and then, as if in doubt on this point, he adds, that it may be the result of the “nature of the tissues and of the surrounding conditions.” He even makes the suggestion, somewhat further on, that the colors may be warning, although it is confessedly unknown that these fish are distasteful to fish-devouring animals.

In amphibians the crest on the back of the male triton, which becomes colored along its edge, is described as a secondary sexual character. The vocal sacs, present in some species of frogs, are found sometimes in both sexes, but more highly developed in the males. In other species, as in the toad, it is the male alone that sings. In the reptiles we find that the two sexes of the turtles are colored alike, and this holds also for the crocodiles. Some male turtles make sounds at the breeding season, and the same is true for the crocodiles, the males of which are said to make a “prodigous display.” In snakes the males are smaller, as a rule, than the females, and the colors are more strongly pronounced, and although some snakes are very brilliantly colored, Darwin puts this down either to protective coloration, or to mimicry of other kinds of snakes. But surely the extremely brilliant colors of many snakes cannot be accounted for in any of these ways. The cause of the color of the venomous kinds, that are supposed to be imitated by the others, “remains to be explained and this may perhaps be sexual selection.”

“It does not, however, follow because snakes have some reasoning power, strong passions and mutual affection, that they should likewise be endowed with sufficient taste to admire brilliant colors in their partners, so as to lead to the adornment of the species through sexual selection. Nevertheless, it is difficult to account in any other manner for the extreme beauty of certain species; for instance, of the coral-snakes of South America, which are of a rich red with black and yellow transverse bands.”

In lizards the erectile crests of the male _Anolis_, the brilliant throat patches of _Sitaria minor_, which is colored blue, black, and red, the skinny appendages present on the throat of the little lizards of the genus Draco, which in the beauty of their colors baffle description, are given as cases of sexual adornment. In the last case cited the ornaments are present, however, in both sexes. The remarkable horns in the males of different species of chameleons are imagined to have been acquired through the battle of the males with each other.