Complex phenotypes can be modeled as networks of component traits connected by genetic, developmental, or functional interactions. Aposematism, which has evolved multiple times in poison frogs (Dendrobatidae), links a warning signal to a chemical defense against predators. Other traits are involved in this complex phenotype. Most aposematic poison frogs are ant specialists, from which they sequester defensive alkaloids. We found that aposematic species have greater aerobic capacity, also related to diet specialization. To characterize the aposematic trait network more fully, we analyzed phylogenetic correlations among its hypothesized components: conspicuousness, chemical defense, diet specialization, body mass, active and resting metabolic rates, and aerobic scope. Conspicuous coloration was correlated with all components except resting metabolism. Structural equation modeling on the basis of trait correlations recovered "aposematism" as one of two latent variables in an integrated phenotypic network, the other being scaling with body mass and physiology ("scale"). Chemical defense and diet specialization were uniquely tied to aposematism whereas conspicuousness was related to scale. The phylogenetic distribution of the aposematic syndrome suggests two scenarios for its evolution: (i) chemical defense and conspicuousness preceded greater aerobic capacity, which supports the increased resource-gathering abilities required of ant-mite diet specialization; and (ii) assuming that prey are patchy, diet specialization and greater aerobic capacity evolved in tandem, and both traits subsequently facilitated the evolution of aposematism.allometry | biodiversity | multivariate | comparative methods S pecies display an array of developmentally complex and recurrent phenotypes produced by common ancestry, natural selection, and genetic drift (1, 2). Phenotypic integration is evidenced by developmental, heritable, and functional relationships among different sets of traits that coevolve and give rise to complex phenotypes (1). One such phenotype is aposematism, defined as the co-occurrence of warning signals and defense mechanisms (3). However, other traits (e.g., diet specialization and gregarious behavior) may also contribute to the aposematic phenotype (4). Here, we model the aposematic syndrome in poison frogs as a phenotypic network integrating several traits: conspicuousness, alkaloid sequestration, diet specialization, body mass, and metabolic rates.A. R. Wallace, who introduced the concept of aposematism, struggled to understand its evolution (5, 6). In aposematism, predators associate conspicuous prey with an unprofitable meal and refrain from attack (7). Thus, defended, the prey may expand its own foraging capabilities (8). The evolution of aposematism is complex for at least four reasons. First, some aposematic organisms synthesize or sequester defensive compounds (9). Second, conspicuousness (high contrast relative to background) signals to the predator that an attack is costly (10). Third, conspicuousness and d...