Drought stress mitigation by nitrogen in boreal forests inferred from stable isotopes.

Forest growth in most parts of the boreal zone is originally limited by low temperatures and low nitrogen availability. Due to the rapid climate warming at high latitudes, an increasing forest area is switching to drought limitation, especially in continental and southern parts of the boreal forest. Studies addressing this issue were mostly dendrochronological and remote-sensing analyses focusing on climatic effects, but not answering the question whether drought is effective alone or in combination with nitrogen shortage at limiting the forests' productivity and vitality. Here we show in a case study from larch forests of Mongolia with a combination of stable isotope analyses, tree-ring analysis and bioindication of the local variability of livestock densities using epiphytic lichens that, in the studied highly drought-prone forests at the southern fringe of the boreal forest in Inner Asia, the trees' vulnerability to drought is modified by nitrogen fertilization from livestock kept in the vicinity and the edge of the forests. The most likely mechanism behind this drought-nitrogen interaction is the reduction of stomatal conductance, which is known to be induced by low nitrogen levels in plants. Nitrogen fertilization by the livestock could, thus, shorten the times of stomatal closure and thereby increase tree growth, which we measured as radial stem increment. Even though the underlying mechanisms, which were so far examined in angiosperms, should be experimentally tested for conifers, our results indicate that focusing on water alone is not enough to understand the climate change response of drought-limited boreal forests.