Evolutionary theory has emphasized that the evolution of single traits cannot be understood in isolation when pleiotropy is present. Widespread pleiotropy causes the appearance of stabilizing selection on metric traits owing to joint effects with fitness, and results in the genetic variation being concentrated in relatively few combinations of the measured traits. In this review, we show how trait combinations with high levels of genetic variation can be used to uncover fitness optima that are defined by apparent stabilizing selection. Defining fitness optima in this way could provide one avenue by which researchers can overcome the problem posed by measuring the myriad of traits that must influence fitness, or by measuring total fitness itself.
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