Fruits, Frugivores, and the Evolution of Phytochemical Diversity

Plants produce an enormous diversity of secondary metabolites, but the evolutionary mechanisms that maintain this diversity are still unclear. The interaction diversity hypothesis suggests that complex chemical phenotypes are maintained because different metabolites benefit plants in different pairwise interactions with a diversity of other organisms. In this synthesis, we extend the interaction diversity hypothesis to consider that fruits, as potential hotspots of interactions with both antagonists and mutualists, are likely important incubators of phytochemical diversity. We provide a case study focused on the Neotropical shrub Piper reticulatum that demonstrates: 1) secondary metabolites in fruits have complex and cascading effects for shaping the outcome of both mutualistic and antagonistic fruit–frugivore interactions, and; 2) fruits can harbor substantially higher levels of phytochemical diversity than leaves, even though leaves have been the primary focus of plant chemical ecology research for decades. We then suggest a number of research priorities for integrating chemical ecology with fruit–frugivore interaction research and make specific, testable predictions for patterns that should emerge if fruit interaction diversity has helped shape phytochemical diversity. Testing these predictions in a range of systems will provide new insight into the mechanisms driving frugivory and seed dispersal and shape an improved, whole-plant perspective on plant chemical trait evolution

Multiscale variability in nutrients and secondary metabolites in a bat‐dispersed neotropical fruit

Abstract Ripe fleshy fruits contain not only nutrients but also a diverse array of secondary metabolites. Nutrients serve as a reward for mutualists, whereas defensive metabolites protect the fruit against pests and predators. The composition of these chemical traits is highly variable, both across different plants and even within repeating structures on the same individual plant. This intraspecific and intraindividual variation has important fitness consequences for both plants and animals, yet patterns of variation and covariation in nutrients and secondary metabolites are not well understood, especially at smaller scales. Here, we investigate the multiscale variation and covariation between nutrients and defensive metabolites in Piper sancti‐felicis ripe fruits. Means and measures of variation of sugars, proteins, phenolics, and alkenylphenols vary greatly among plants, and at least 50% of the trait variation occurs at the intraindividual level. Also, we found that proteins, but not sugars, were correlated with phenolics and alkenylphenols at multiple scales, suggesting trait variation in protein content may be more constrained than sugars. Our findings emphasize the importance of examining patterns across scales and provide the groundwork to better understand how complex patterns of variation and covariation in nutrients and defensive metabolites shape ecological interactions surrounding fruits