Joint evolution of seed traits along an aridity gradient: Seed size and dormancy are not two substitutable evolutionary traits in temporally heterogeneous environment

Seed size and dormancy are reproductive traits that interact as adaptations to environmental conditions. Here, we explore the evolution of these traits in environments that differ in overall mean favorability and in the extent of temporal predictability. Our model simulates a population of annual plants living in a range of environments that differ in aridity, namely mean annual precipitation and inter-annual variation of this mean precipitation. The optimal fitness curve is investigated assuming density dependence, three alternative hypothetical relationships between seed mass and seed survival in the soil (negative, positive, and independent of mass), and three alternative relationships between survival in soil and precipitation (strong and intermediate negative relationships, and no relationship). Our results show that seed size and dormancy are not two substitutable evolutionary traits; that specific combinations of these two traits are selected in environments that differ in favorability and temporal predictability; that a certain degree of seed dormancy is advantageous not only in temporally unpredictable environments but also in temporally predictable environments with high competition; and that more than one combination of seed size and dormancy (defined in terms of germination fraction) can be optimal, even in spatially homogeneous environments, potentially allowing selection for more variation in these traits within and among species.