Forb Diversity Globally is Harmed by Nutrient Enrichment but Can Be Rescued by Large Mammalian Herbivory

Forbs (“wildflowers”) are important contributors to grassland biodiversity and services, but they are vulnerable to environmental changes that affect their coexistence with grasses. In a factorial experiment at 94 sites on 6 continents, we tested the global generality of several broad predictions arising from previous studies: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment, which benefits grasses at the expense of forbs. (2) Forb cover and richness increase under herbivory by large mammals, especially when nutrients are enriched. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. We found strong evidence for the first, partial support for the second, and no support for the third prediction. Forb richness and cover are reduced by nutrient addition, with nitrogen having the greatest effect; forb cover is enhanced by large mammal herbivory, although only under conditions of nutrient enrichment and high herbivore intensity; and forb richness is lower in more arid sites, but is not affected by consistent climate-nutrient or climate-herbivory interactions. We also found that nitrogen enrichment disproportionately affects forbs in certain families (Asteraceae, Fabaceae). Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs and the ecosystem services they support, but grazing under high herbivore intensity can offset these nutrient effects.