Correlated evolution of seed mass and genome size varies among life forms in flowering plants

Seeds show important variation as plant regenerative units among species, but their evolutionary co-variations with other plant characteristics are still poorly understood. Whilst a positive association of seed mass with genome size (GS) and life forms has already been documented, a broad-scale quantification of their evolutionary correlation and adaptive selection has never been conducted. Here, we tested for correlated evolution of seed mass and GS towards distinct selective regimes related to life form in angiosperms. In particular, we tested the hypothesis that the selection toward lighter seeds and smaller genomes is stronger for annual plants, ensuring high regenerative potential. Using multivariate evolutionary models over a dataset containing 3242 species, we showed an overall positive correlated evolution of seed mass and GS deviating from a pure drift process. Instead, evolutionary changes in seed and genome sizes were driven by adaptive selection towards optimal values differing among life forms. Specifically, the evolutionary optima towards which the seed and genome sizes evolve show a covariation toward small values in annuals, intermediate values in perennial herbs and a trade-off between seed mass and GS in woody plants. Moreover, the evolutionary correlation between seed mass and GS is strongest in annuals as an adaption to complete their life cycle in a short time window, when environmental conditions are favourable for regeneration and development to maturity. The asymmetry in the correlated evolution acting on seed and genome sizes due to life form could explain how life-history traits interplay with functional traits and how plants have evolved diverse successful life-history strategies.