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Evolution__3rd_Edition

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328 PART 3 / Adaptation and Natural Selection<br />

The peacock’s tail is an example of<br />

a costly sexual character<br />

Sexual selection works mainly<br />

by...<br />

. . . male competition ...<br />

. . . and female choice<br />

these characters exist. If a population contains some types with higher survival than<br />

other types, natural selection will fix the former and eliminate the latter.<br />

Characters that reduce survival can be called “deleterious” or “costly.” One large<br />

class of apparently costly characters are those found usually only in males and which<br />

Darwin called secondary sexual characters. The primary sexual characters are things<br />

like genitalia that are needed for breeding. The secondary sexual characters are not<br />

actually needed for breeding, but they function during reproduction. The peacock’s<br />

“tail” (or, more exactly, train) is an example. In many other bird species too, the males<br />

have tails or other extravagantly developed and brightly colored structures. A peacock<br />

could inseminate a female just as well without his remarkable tail, and in that sense it is<br />

a secondary, not a primary, sexual organ. The peacock’s tail almost certainly reduces<br />

the male’s survival (though this disadvantage has never actually been demonstrated) as<br />

the tail reduces maneuverability, powers of flight, and makes the bird more conspicuous;<br />

its growth must also impose an energetic cost. Why are these costly characters not<br />

eliminated by selection?<br />

12.4.2 Sexual selection acts by male competition and female choice<br />

Darwin’s solution was his theory of sexual selection. He defined the process by saying<br />

that it “depends on the advantage which certain individuals have over other individuals<br />

of the same sex and species, in exclusive relation to reproduction.” A structure produced<br />

by sexual selection in males exists not because of the struggle for existence, but<br />

because it gives the males that possess it an advantage over other males in the competition<br />

for mates. Darwin’s idea is that the reduced survival of peacocks with long, colorful<br />

tails is more than compensated by their increased “advantage in reproduction.”<br />

Darwin discussed two kinds of sexual selection. One is for males to compete among<br />

each other for access to females. Male competition can take the form of direct fighting,<br />

or it can be more subtle. Some male insects, for instance, can remove sperm from<br />

females they are copulating with a sperm that was stored from matings with previous<br />

males. However, we shall not discuss adaptations of sperm competition here, or other<br />

adaptations of male competition, because they do not pose deep theoretical questions.<br />

The situation is different for Darwin’s other mechanism: female choice.<br />

A structure like the peacock’s tail cannot plausibly be explained by male competition.<br />

It would be no use in fighting a indeed it would reduce the male’s fighting power<br />

a and no one has ever thought up a more subtle competitive function for the tail.<br />

Darwin suggested that the tail exists instead because females preferentially mate with<br />

males that have longer, brighter, or more beautiful tails. If they do, the mating advantage<br />

of males with longer tails will compensate a corresponding amount of reduced male<br />

survival.<br />

Darwin’s main argument for the importance of sexual selection was comparative.<br />

Sexual selection should operate more powerfully in polygamous than in monogamous<br />

species. In a polygynous species, in which several females mate with one male (and<br />

other males do not breed at all), a single male can potentially breed with more females<br />

than under monogamy; selection in favor of adaptations that enable males to gain<br />

access to females (whether by male competition or female choice) is proportionally<br />

..

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