Loss of taxonomic and functional diversity and decrease in primary productivity with nitrogen enrichment after a long-term release from grazing in an alpine grassland

Abstract

Moderate grazing can sustain high species diversity and productivity. However, nitrogen enrichment often reduces species richness while promoting primary productivity, which contradicts the traditional understanding of the positive effect of plant diversity on productivity. Whether the responses of diversity and productivity to N enrichment on a long-term scale conform to those on short-term scale. Furthermore, chemical N forms may alter species richness or functional diversity because of species preferences for different N forms. We used data from a 19-year field experiment with manipulated livestock exclusion and the addition of N in three chemical forms (ammonium, nitrate, and their mixture) in an alpine grassland to test their effects on diversity and aboveground productivity. Productivity in no N without grazing plots (Ctr) initially increased by 66 % during the first five years after livestock exclusion (0.53 kg m^-2), following which productivity fluctuated at a similar level as that in winter livestock grazing (0.32 kg m^-2) for the next eight years and ultimately decreased to the level of 0.24 kg m^-2, below that in Gr with continued species loss. The addition of N forms in grazing exclusion increased productivity, accompanied by the loss of rare species in the first five years. Afterward, productivity decreased, and it took longer than the control to reach the same level as that seen in winter grazing. Then, productivity continued to decline with more species loss. The functional diversity of leaf traits and plant growth forms decreased. Ammonium favored grasses with guerrilla and phalanx rhizomes, nitrate favored sedges and forbs with guerrilla rhizomes, and ammonium-nitrate sustained a relatively higher diversity of plant growth forms. Our general conclusion is that grasslands with moderate grazing are beneficial for sustaining diverse species and high productivity. N enrichment reduces plant diversity, and the increase in productivity driven by N is only transitional, as productivity declines in the long run, with continuous N addition.

Keywords: Functional diversity; Livestock grazing; Nitrogen addition; Primary productivity; Species richness.