The many dimensions of phytochemical diversity: linking theory to practice.

Research on the ecological and evolutionary roles of phytochemicals has recently progressed from studying single compounds to examining chemical diversity itself. A key conceptual advance enabling this progression is the use of species diversity metrics for quantifying phytochemical diversity. In this perspective, we extend the theory developed for species diversity to further our understanding of what exactly phytochemical diversity is and how its many dimensions impact ecological and evolutionary processes. First, we discuss the major dimensions of phytochemical diversity - richness, evenness, functional diversity, and alpha, gamma and beta diversity. We describe their potential independent roles in biotic interactions and the practical challenges associated with their analysis. Second, we re-analyse the published and unpublished datasets to reveal that the phytochemical diversity experienced by an organism (or observed by a researcher) depends strongly on the scale of the interaction and the total amount of phytochemicals involved. We argue that we must account for these frames of reference to meaningfully understand diversity. Moving from a general notion of phytochemical diversity as a single measure to a precise definition of its multidimensional and multiscale nature yields overlooked testable predictions that will facilitate novel insights about the evolutionary ecology of plant biotic interactions.