The antics of male birds at the mating season, their courtship so-called, has played an important rôle in Darwin’s theory of sexual selection. The behavior of many birds at this time is of such a kind as to suggest that the male is exhibiting his plumage before the female for the “purpose” of influencing her choice. The whole paraphernalia of human psychology is imported into the situation and both the consciousness of the male, his intentions so to speak, and the supposed esthetic response or choice of the female is invoked. Even though it be granted that the words that we must make use of, borrowed from human behavior, are such as to imply much more in the direction of consciousness and purpose than is desirable, and that most of the behavior of animals should be stated in a more roundabout and objective way, yet the theory will only work out on the assumption that the female chooses in some sense the more brilliant or ornamental (or effective) male, whether she is “conscious” or unconscious of intention. I doubt if anyone to-day would care to defend seriously the theory on the grounds of consciousness or esthetic value of the exhibition, despite the fact that Darwin’s language often takes this turn and the less-guarded statements of some of his disciples, such as Romanes, show little hesitation in anthropo-morphologizing the entire situation. It is, however, not necessary for the working out of the theory that this complication be introduced into it, for if the female is more likely to mate with a more brilliantly colored than a less brilliantly colored male, the theory may be made to apply regardless of whether she is “conscious” or not of the difference to which she responds.
But there are weighty arguments against such an interpretation of the behavior of the male and female during courtship. In the first place, there is almost no direct evidence to show that the female mates with the more ornamental male. As this is the all-essential requirement of the theory, the almost complete absence of facts in its support leaves the theory resting on a theoretical assumption. It can scarcely pass unnoticed that while there exists a large mass of data describing the secondary sexual characters, there is practically nothing in this accumulation to show that the female makes her selection on differences in coloration or ornamentation. And on the other hand, there is some evidence showing that the female is ready to succumb to the aggressiveness of the male rather than that she “chooses” him.
The behavior of the male under sexual excitement is often described to be of a kind to exhibit before the female his peculiar adornments. That the “purpose” of his exhibition is to show himself off before the female may be conceded, with reservations as to what is meant here by “purpose.” That the male is conscious of the probable results of his conduct is scarcely probable the first time he courted; but that he may have found out the most probable result after the first attempt through “associative memory” is in accord with what the study of “animal behavior” has shown to be possible. In this sense purpose would mean a line of conduct that experience had shown to lead to a certain end. Anticipation or far-sightedness would henceforth characterize such a reaction. Here, however, we venture on very dubious grounds. But the display of the male may be purposeful in a much simpler sense. His activity may be an inborn reflex to visual or other sensory stimuli that is a part of his attack on the female, or possibly a series of reflexes that we may register under the old unanalyzed terms of “desire and fear.” The action calls forth a responsive reflex in the female, for the sexual act is not entirely active on one side, passive on the other, but consists of a series of interreactions on the part of each sex, which, if they pursue a given course, leads to the final mating. The mutual responses appear to follow an automatic course in many cases if the individuals are sexually ready to mate and the environment is propitious. Types of behavior of this kind must be familiar to anyone who has observed domesticated and semi-domesticated animals. The purpose of the display may mean no more than a reaction that leads to a result propitious to the perpetuation of the species if the situation is ripe for such an outcome.
This conclusion still leaves open the question as to whether the display is more likely to be successful, if certain special characters possessed by the species are exhibited. In the absence of any sufficient evidence to show that this is so, and in the light of the very great danger of projecting “our human standards” into the world of other animals, and in view of the fact that related species without such marks are as successful in maintaining themselves, I can not but think that at present we have a good deal to lose in the way of scientific procedure and nothing to gain of scientific value in accepting Darwin’s interpretation of sexual selection based on the display of the male as furnishing an opportunity to the female to make the “best” selection amongst her suitors on the basis of his adornment.
An excellent opportunity to study the problem as to “choosing” by the female is furnished by the mutant races of Drosophila, some of which, differing in a single mutant gene, have wings as different in coloration as black, yellow, or gray, and eyes as differently colored as white, vermilion, or red. Sturtevant put a yellow female with a gray (wild-type) male and a yellow male. The male that first mated was noted and the trio discarded. The female “chose” the gray males 25 times and the yellow only 8 times. In the control combination, where a gray female “chose” between the same two kinds of males, she took the gray male 60 times and the yellow male 12 times. In both cases it “appears” that the female “prefers” the gray male, but this deduction may give an entirely wrong impression as to what is taking place, for the result would be the same in kind if the gray male were more active and mated quicker. This was tested by putting a gray and a yellow female with a gray male and then for control a gray and a yellow female with a yellow male. The result was as follows:
| Red ♂ | {Gray ♀ | 25 | Yellow ♂ | {Gray ♀ | 12 | |
| {Yellow ♀ | 31 | {Yellow ♀ | 30 |
Here the gray male mated slightly oftener with the yellow female than with the other, whereas the yellow male mated much oftener with the yellow female than with the gray one. Both results are explicable on the view that the yellow female, being less active, is more easily captured by the yellow male than is the gray female. This view fits in also with the former experiment, where the yellow male is much less successful than the more active gray male. Such a conclusion gives a more consistent explanation of all the facts than does the theory of female choice, for on the latter we must suppose that the yellow females prefer the gray males and the yellow male prefers the yellow females, etc.
The following results were obtained by Sturtevant when red and white eyed flies were competing:
| Red ♂ | {Red ♀ | 54 | Red ♀ | {Red ♂ | 53 | |
| {White ♀ | 82 | {White ♂ | 14 | |||
| White ♂ | {Red ♀ | 40 | White ♀ | {Red ♂ | 62 | |
| {White ♀ | 93 | {White ♂ | 19 |
The outcome can be interpreted in the same way as the yellow-gray competition. The red male wins by virtue of his greater activity, while the white female is chosen more often, especially by the white male, because of her passivity (or weaker resistance). It may be claimed that these results do not show that the female does not choose, for such choice, if made, would be swamped by another condition of the experiment, viz, the greater aggressiveness of one kind of male and greater passivity of the other kind of female. This, of course, is true, but the experiment still shows that in these flies other influences are so much greater than “choice” by the female, if it exists, that the postulated effect of the latter practically disappears from the situation.