Spatially divergent trends of nitrogen versus phosphorus limitation across European forests

Nitrogen (N) and phosphorus (P) are essential nutrients that widely limit plant growth in global terrestrial ecosystems. Rising atmospheric CO₂ concentration generally stimulates terrestrial net primary productivity and consequently may cause or aggravate N and P limitation due to a dilution effect, but the spatial variation of temporal trends in N versus P limitation and its key regulating factors is poorly understood. Using the leaf N:P ratio of 15 dominant tree species as an indicator, we analysed the spatial variation of plot-level shift towards N or P limitation across 163 European forest plots during 1995-2017. Phosphorus limitation increased from 25% to 33% of the studied plots between 1995-1997 and 2015-2017, while N limitation occurred in a negligible number of plots. A major proportion (56%) of the plots showed no significant trend in leaf N:P ratio, implying no shifts in N versus P limitation status. In the remaining plots, 38% of the plots showed a significant increase of leaf N:P ratio and only 6% of the plots showed a significant decrease of leaf N:P ratio. The spatial variation in the rate of decrease in leaf N:P ratio was associated with a significant decrease in leaf N concentration and mainly explained by the rate of decrease in N deposition. In contrast, the spatial variation in the rate of increase in leaf N:P ratio was associated with a significant decrease in leaf P concentration and mainly explained by forest category (broadleaf vs. conifer), mean annual temperature and soil C:N ratio. Our findings highlight a remarkable spatial divergence in temporal trends of nutrient limitation status across European forests over the past two decades, but overall, P is becoming more limiting versus N, especially in broadleaved forests.