Phenotypic Diversity Arises from Secondary Signal Loss in the Elaborate Visual Displays of Toucans and Barbets.

Complexity and diversity are fundamental characteristics of life, but the relationship between the two remains murky. For example, both gaining and losing complexity can support diversity-so how exactly does complexity influence the emergence of unique phenotypes? Here we address this question by examining how complexity underlies the diversity of elaborate visual displays in the avian clade Ramphastides (toucans and barbets). These species communicate in part by using body movement and colorful ornaments on the tail. We find that sexual size dimorphism predicts the evolution of one specific signal, the tail-cock gesture, implying that tail cocking is more likely to evolve under stronger sexual selection. We also discover process-level constraints on the evolution of complexity: signals are gained along a strict order of operations, where the tail-cock gesture arises before other colors and gestures. Yet virtually any signal can be lost at any time. As a result, many extant phenotypes were more likely to arise through loss of complexity, highlighting the importance of secondary signal loss to phenotypic diversity. Collectively, our results demonstrate how sexual selection catalyzes the evolution of complex phenotypes, which indirectly support diversity by allowing different traits to be modified or lost in the future.