Physiological and morphological responses to shade and nutrient additions of Claytonia virginica (Portulacaceae): implications for the "vernal dam" hypothesis

Abstract

Because of their unique phenology and physiology, spring ephemeral herbs are believed to play an important role in intrasystem nutrient cycling in deciduous forest ecosystems. It was hypothesized that they function as a "vernal dam" by temporarily sequestering nutrients and preventing leaching from the system during a period of high nutrient availability. However, spring ephemerals require high-irradiance growing conditions. How do their physiological and morphological responses to ambient light and shade limit their ability to sequester excess nutrients? We performed field experiments using Claytonia virginica L. as a model to test several responses to shade and increasing levels of nutrient additions. We also examined the biomass responses and nutrient storage capacities of other spring ephemeral herbs. In C. virginica, shading reduced ribulose 1,5-bisphosphate carboxylase-oxygenase (Rubisco) activity, photosynthesis rate, specific leaf weight, leaf width/length (W/L), and biomass; nutrient additions increased W/L and biomass only under unshaded conditions. Other herbs responded similarly but reached maximum biomass at lower nutrient addition levels than C. virginica. Shading reduced and nutrient additions increased nitrogen and phosphorus concentrations in both C. virginica and other herbs. Shaded herbs generally reached nutrient saturation at lower nutrient addition levels than unshaded herbs. Overall, unshaded plants sequestered larger amounts of nutrients than shaded plants. This pattern is best explained by a reduction in biomass under shaded conditions. We concluded that C. virginica and other spring herbs, although important components in forest nutrient cycling in the early spring, are limited in their capacity to store excess nutrients, particularly when shaded.Key words: Claytonia virginica, nutrient cycling, spring ephemerals, vernal dam.