Chasing the fitness optimum: temporal variation in the genetic and environmental expression of life-history traits for a perennial plant

Abstract The ability of plants to track shifting fitness optima is crucial within the context of global change, where increasing environmental extremes may have dramatic consequences to life history, fitness, and ultimately species persistence. However, to track changing conditions relies upon the complex relationship between genetic and environmental variance, where selection may favor plasticity, the evolution of genetic differences, or both depending on the spatial and temporal scale of environmental heterogeneity. Over three years, we compared the genetic and environmental components of phenological and life-history variation in a common environment for the spring perennial Geum triflorum. Populations were sourced from alvar habitats that exhibit extreme, but predictable annual flood-desiccation cycles and prairie habitats that exhibit similar, but less predictable variation in water availability. Narrow-sense heritabilities were generally higher for early life history (emergence probability) relative to later life history traits (total seed mass), indicating that traits associated with establishment within an environment are under stronger genetic control relative to later life-history fitness expressions, where plasticity may play a larger role. This pattern was particularly notable in seeds sourced from environmentally extreme, but predictable alvar habitats relative to less predictable prairie seed sources. Fitness landscapes based on seed source origin, largely characterized by varying water availability and flower production, described selection as the degree of maladaptation to the prairie common garden environment relative to seed source environment. Plants from alvar populations were consistently closer to the fitness optimum across all years. Annually, the breadth of the fitness optimum expanded primarily along a moisture gradient, with inclusion of more populations onto the expanding optimum. These results highlight the importance of temporally and spatially varying selection for the evolution of life history, indicating plasticity within perennial systems may over time become the primary mechanism to track fitness for later life history events.