The Effects of Temporal Variation on Fitness, Functional Traits, and Species Distribution Patterns.

Temporal variation is a powerful source of selection on life history strategies and functional traits in natural populations. Theory predicts that the rate and predictability of fluctuations should favor distinct strategies, ranging from phenotypic plasticity to bet-hedging, which are likely to have important consequences for species distribution patterns and their responses to environmental change. To date, we have few empirical studies that test those predictions in natural systems, and little is known about how genetic, environmental, and developmental factors interact to define the "fluctuation niche" of species in temporally variable environments. In this study, we evaluated the effects of hydrological variability on fitness and functional trait variation in three closely related plant species in the genus Lasthenia that occupy different microhabitats within vernal pool landscapes. Using a controlled greenhouse experiment, we manipulated the mean and variability in hydrological conditions by growing plants at different depths with respect to a shared water table and manipulating the magnitude of stochastic fluctuations in the water table over time. We found that all species had similarly high relative fitness above the water table, but differed in their sensitivities to water table fluctuations. Specifically, the two species from vernal pools basins, where soil moisture is controlled by a perched water table, were negatively affected by the stochasticity treatments. In contrast, a species from the upland habitat surrounding vernal pools, where stochastic precipitation events control soil moisture variation, was insensitive to experimental fluctuations in the water table. We found strong signatures of genetic, environmental (plastic), and developmental variation in four traits that can influence plant hydrological responses. Three of these traits varied across plant development and among experimental treatments in directions that aligned with constitutive differences among species, suggesting that multiple sources of variation align to facilitate phenotypic matching with the hydrological environment in Lasthenia. We found little evidence for predicted patterns of phenotypic plasticity and bet-hedging in species and traits from predictable and stochastic environments, respectively. We propose that selection for developmental shifts in the hydrological traits of Lasthenia species has reduced or modified selection for plasticity at any given stage of development. Collectively, these results suggest that variation in species' sensitivities to hydrological stochasticity may explain why vernal pool Lasthenia species do not occur in upland habitat, and that all three species integrate genetic, environmental, and developmental information to manage the unique patterns of temporal hydrological variation in their respective microhabitats.