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Over the years, biologists have suggested two main pathways by which sexual selection may have shaped the evolution of male birdsong. In the first, male competition and intrasexual selection produce relatively short, simple songs used mainly in territorial behavior. In the second, female choice and intersexual selection produce longer, more complicated songs used mainly in mate attraction; like such visual ornamentation as the peacock's tail, elabo-rate vocal characteristics increase the male's chances of being chosen as a mate, and he thus enjoys more repro-ductive success than his less ostentatious rivals. The two pathways are not mutually exclusive, and we can expect to find examples that reflect their interaction. Teasing them apart has been an important challenge to evolutionary biol-ogists.


Early research confirmed the role of intrasexual selection.In a variety of experiments in the field, males responded aggressively to recorded songs by exhibiting territorial behavior near the speakers. The breakthrough for research into intersexual selection came in the development of a newtechnique for investigating female response in the labor-atory. When female cowbirds raised in isolation in sound-proof chambers were exposed to recordings of male song,they responded by exhibiting mating behavior. By quanti-fying the responses, researchers were able to determine what particular features of the song were most important. In further experiments on song sparrows, researchers found that when exposed to a single song type repeated several times or to a repertoire of different song types, females responded more to the latter. The beauty of the experi-mental design is that it effectively rules out confounding variables; acoustic isolation assures that the female can respond only to the song structure itself. 


If intersexual selection operates as theorized, males with more complicated songs should not only attract females more readily but should also enjoy greater reproductive success. At first, however, researchers doing fieldwork with song sparrows found no correlation between larger reper-toires and early mating, which has been shown to be one indicator of reproductive success; further, common measuresof male quality used to predict reproductive success, such as weight, size, age, and territory, also failed to correlate with song complexity. 


The confirmation researchers had been seeking was finally achieved in studies involving two varieties of war-blers. Unlike the song sparrow, which repeats one of its several song types in bouts before switching to another, the warbler continuously composes much longer and more vari-able songs without repetition. For the first time, researchers found a significant correlation between repertoire size and early mating, and they discovered further that repertoire size had a more significant effect than any other measure of male quality on the number of young produced. The evi-dence suggests that warblers use their extremely elaborate songs primarily to attract females, clearly confirming the effect of intersexual selection on the evolution of birdsong.

The passage suggests that the song sparrow experiments mentioned in lines 37-43 failed to confirm the role of intersexnal selection because

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