Global-scale patterns of nutrient density and partitioning in forests in relation to climate

Knowledge of nutrient storage and partitioning in forests is imperative for ecosystem models and ecological theory. Whether the nutrients (N, P, K, Ca, and Mg) stored in forest biomass and their partitioning patterns vary systematically across climatic gradients remains unknown. Here, we explored the global-scale patterns of nutrient density and partitioning using a newly compiled dataset including 372 forest stands. We found that temperature and precipitation were key factors driving the nutrients stored in living biomass of forests at global scale. The N, K, and Mg stored in living biomass tended to be greater in increasingly warm climates. The mean biomass N density was 577.0, 530.4, 513.2, and 336.7 kg/ha for tropical, subtropical, temperate, and boreal forests, respectively. Around 76% of the variation in biomass N density could be accounted by the empirical model combining biomass density, phylogeny (i.e., angiosperm, gymnosperm), and the interaction of mean annual temperature and precipitation. Climate, stand age, and biomass density significantly affected nutrients partitioning at forest community level. The fractional distribution of nutrients to roots decreased significantly with temperature, suggesting that forests in cold climates allocate greater nutrients to roots. Gymnosperm forests tended to allocate more nutrients to leaves as compared with angiosperm forests, whereas the angiosperm forests distributed more nutrients in stems. The nutrient-based Root:Shoot ratios (R:S), averaged 0.30 for R:S_N , 0.36 for R:S_P , 0.32 for R:S_K , 0.27 for R:S_Ca , and 0.35 for R:S_Mg , respectively. The scaling exponents of the relationships describing root nutrients as a function of shoot nutrients were more than 1.0, suggesting that as nutrient allocated to shoot increases, nutrient allocated to roots increases faster than linearly with nutrient in shoot. Soil type significantly affected the total N, P, K, Ca, and Mg stored in living biomass of forests, and the Acrisols group displayed the lowest P, K, Ca, and Mg.