Phenotypic diversity acts as a higher-order functional trait of populations

Functional traits are usually assigned to individuals, yet some properties exist only as differences among population members. Whether such higher-order properties predict collective performance remains unclear. Here we used the aquatic plant Spirodela polyrhiza to test whether phenotypic and genetic differences among genotypes predict non-additive population growth. We grew twelve Japanese strains alone and in all pairwise mixtures and quantified their growth dynamics. Mixtures sometimes outperformed the mean of their component monocultures, and some exceeded the better-performing monoculture, although diversity effects were negative on average. Positive effects were most strongly associated with between-strain differences in frond colour and morphology, identifying specific dimensions of phenotypic diversity as population-level predictors of synergistic growth. Genome-wide higher-level association mapping identified loci where between-strain diversity was associated with growth effects, implicating stress responses, secondary metabolism and frond morphology. Loci associated with reduced performance implicated chemical perception and self/non-self recognition, whereas those associated with positive effects reflected functional differentiation in stress responses and physiological niches. Our findings suggest that some traits generate new functions at the population level through variation among individuals, making diversity itself a higher-order functional property.