Aggregation Pheromone for an Invasive Mussel Consists of a Precise Combination of Three Common Purines

Most marine benthic invertebrates have a pelagic larval phase, after which they settle preferentially on or near conspecific adults, forming aggregations. Although settlement pheromones from conspecific adults have been implicated as critical drivers of aggregation for more than 30 years, surprisingly few have been unambiguously identified. Here we show that in the invasive dreissenid mussel Mytilopsis sallei (an ecological and economic pest), three common purines (adenosine, inosine, and hypoxanthine) released from adults in a synergistic and precise ratio (1:1.125:3.25) serve as an aggregation pheromone by inducing conspecific larval settlement and metamorphosis. Our results demonstrate that simple common metabolites can function as species-specific pheromones when present in precise combinations. This study provides important insights into our understanding of the ecology and communication processes of invasive organisms and indicates that the combination and ratio of purines might be critical for purine-based signaling systems that are fundamental and widespread in nature.