Leaf P increase outpaces leaf N in an Inner Mongolia grassland over 27 years.

The dynamics of leaf nitrogen (N) and phosphorus (P) have been intensively explored in short-term experiments, but rarely at longer timescales. Here, we investigated leaf N : P stoichiometry over a 27-year interval in an Inner Mongolia grassland by comparing leaf N : P concentration of 2006 with that of 1979. Across 80 species, both leaf N and P increased, but the increase in leaf N lagged behind that of leaf P, leading to a significant decrease in the N : P ratio. These changes in leaf N : P stoichiometry varied among functional groups. For leaf N, grasses increased, woody species tended to increase, whereas forbs showed no change. Unlike leaf N, leaf P of grasses and forbs increased, whereas woody species showed no change. Such changes may reflect N deposition and P release induced by soil acidification over the past decades. The interannual effect of precipitation may somewhat have reduced the soil available N, leading to the more modest increase of leaf N than of leaf P. Thus, leaf N : P stoichiometry significantly responded to long-term environmental changes in this temperate steppe, but different functional groups responded differently. Our results indicate that conclusions of plant stoichiometry under short-term N fertilization should be treated with caution when extrapolating to longer timescales.

Leaf nitrogen:phosphorus stoichiometry across Chinese grassland biomes.

Leaf N and P stoichiometry covaries with many aspects of plant biology, yet the drivers of this trait at biogeographic scales remain uncertain. Recently we reported the patterns of leaf C and N based on systematic census of 213 species over 199 research sites in the grassland biomes of China. With the expanded analysis of leaf P, here we report patterns of leaf P and N:P ratios, and analyze the relative contribution of climatic variables and phylogeny in structuring patterns of leaf N:P stoichiometry. Average values of leaf P and N:P ratio were 1.9 mg g(-1) and 15.3 (mass ratio), respectively, consistent with the previous observation of a higher N:P ratio in China's flora than the global averages (ca. 13.8), resulting from a lower leaf P. Climatic variables had very little direct correlation with leaf P and N:P ratios, with growing season precipitation and temperature together explaining less than 2% of the variation, while inter-site differences and within-site phylogenetic variation explained 55 and 26% of the total variation in leaf P and N:P ratios. Across all sites and species, leaf N and P were highly positively correlated at all levels. However, the within-site, within-species covariations of leaf N and P were weaker than those across sites and across species. Leaf N and P relationships are driven by both variation between sites at the landscape scale (explaining 58% of the variance) and within sites at the local scale (explaining 24%), while the climatic factors exerted limited influence (explaining less than 3%). In addition, leaf N:P ratios in two dominant genera Kobresia and Stipa had different responses to precipitation. This study suggests that geographic variation and between-species variation, rather than climatic variation, are the major determinants of grassland foliar stoichiometry at the biome level.