Fluctuating patch boundaries in a native annual forb: the roles of niche and dispersal limitation

Abiotic, biotic, and dispersal constraints jointly control the spatial distributions of species, but few studies have directly evaluated how these forces interact and vary over time to create dynamic spatial distributions. Through three years of observation and two years of field manipulation, I investigated simultaneous constraints on the spatial distribution of Lupinus nanus, a native annual legume that grows in dense patches in California grasslands. I transplanted L. nanus across its own patch boundaries, yet within apparently suitable habitat, and assessed the demographic success of naturally occurring and seeded plants in plots with and without competitor removal in years that varied in temperature and rainfall. Core sites were defined as those consistently densely occupied, whereas peripheral sites were densely occupied only during some years, and exterior sites were consistently unoccupied or very sparsely occupied. Site types (core, periphery, and exterior) differed in soil moisture, P, and NO3. Competition limited emergence in all site types in the dry/warm year and in patch peripheries in the wet/cool year. Population fitness (seeds produced per seed added) was >1.0 in cores during all years. Peripheral sites had fitness near replacement in the wet/cool year, which was greatly increased by competition removal. Exterior fitness was <1.0 in both experimental years, regardless of seed addition and competitor removal. Seed addition did not increase site-specific fitness, and a seed bank was found to be present in all site types. Herbivory was greater in patch cores and peripheries than in exteriors. Soil variation exerted the most consistent control over patch limits, while competition played an intermittent role in excluding Lupinus from patch peripheries. The dynamic distribution of L. nanus is the product of temporal variation in specific abiotic and biotic niche axes, primarily soil characteristics and competition, rather than dispersal limitation.