Non-interacting impacts of fertilization and habitat area on plant diversity via contrasting assembly mechanisms

Diversity and Distributions



The local- and regional-based forms of anthropogenic change reducing grassland diversity are generally identified, but these scale-dependent processes tend to co-occur with unclear interactive effects. Here, we explicitly test how common local and regional perturbations simultaneously affect plant alpha and beta diversity in a multiyear community assembly experiment using fragments of grassland habitat of various sizes. We hypothesized that local disturbances and decreasing patch size would interact, suppressing local diversity while homogenizing composition among patches.


North America.


We conducted a three-year grassland assembly experiment, factorially manipulating local perturbation (nitrogen addition and mowing) and patch area for 36 patches over 13 ha. We quantified the individual and interactive effects of these local and regional factors on plant alpha and beta diversity within (quadrat scale) and among patches (patch scale). We also used a null model approach to disentangle between stochastic- and niche-based assembly mechanisms.


We detected a gradient of assembly outcomes driven by two non-interacting factors—the effects of N fertilization on alpha (negative) and beta (positive) diversity regardless of spatial scale and the scale-dependant effect of increasing patch size on alpha (positive) and beta (positive) diversity. These effects unfolded over time, with the constraints on richness and composition shifting from dispersal-based during the first sampling year to perturbation-and size-based factors at year two and three. Fertilization effects were driven by a mixture of deterministic (i.e., selection at the species level) and stochastic (i.e., random extinctions) processes resulting in a decline in local richness but an increase in spatial heterogeneity in species composition. Area appeared to influence alpha diversity mainly via stochastic “sampling effect”—larger patches represented a larger sample of the regional pool. Niche-based processes, however, led to convergence in beta diversity among smaller patches driving a positive overall effect of area on beta diversity.

Main conclusion

Our results illustrate how diversity regulation in contemporary grasslands can be simultaneously shaped by local and regional factors acting additively but via contrasting assembly mechanisms that operate at different spatial and temporal scales.