Strength of natural or sexual selection FIGURE 12.1 Complex traits, such as behavior and performance, comprise hierarchical suites of interacting subordinate traits at lower levels of biological organization. In general, selection is thought to act most strongly on phenotypic variation in traits at higher levels of biological organization, such as behavior (see also Rhodes and Kawecki this volume) and performance, as indicated by the width of the inverted triangle (and the direction of the arrow). In other words, behavior and performance have strong effects on major components of Darwinian fitness, such as survivorship and fecundity. Lower-level traits include a wide range of suborganismal morphological, physiological, and biochemical phenotypes. These lowerlevel traits, directly and indirectly, influence aspects of whole-organism performance ability that are crucial for survival and reproduction.