No effect of varying soil resource heterogeneity on plant species richness in a low fertility grassland

1. Coexistence theory predicts that greater heterogeneity of resources or other fitness-constraining environmental factors will promote species diversity, yet this classic mechanism of coexistence has rarely been tested in manipulative field experiments. 2. Here we present results from the fourth year of a long-term experiment designed to test the heterogeneity-diversity hypothesis in a low productivity grassland where we directly manipulated the spatial heterogeneity of soil nutrients. In addition to unfertilized controls, we created heterogeneous and homogeneous nutrient enrichment treatments by applying fertilizer to field plots in a patchy or uniform manner. Fertilization treatments were crossed with seed addition and cover reduction treatments, to allow examination of the effects of recruitment limitation and competition from established vegetation on diversity responses to nutrient heterogeneity. 3. In contrast to predictions, we found that spatially heterogeneous soil nutrient supply did not promote increased species richness or greater species sorting relative to a uniform supply of soil nutrients. Instead, both nutrient enrichment treatments depressed plant species richness (especially at small spatial scales) and, depending on cover reduction and on spatial location in the field, increased the abundance of two spreading clonal species. 4. Our results do not support the heterogeneity-diversity hypothesis, and suggest that the response of clonal species to nutrient enrichment may constrain how plant diversity responds to resource heterogeneity. Future tests of the heterogeneity-diversity hypothesis should explicitly consider the scale of resource heterogeneity and foraging capability of the plant species in the community.