Deposited pollen from dominant tree genera drives nutrient composition of throughfall in European forests

Atmospheric deposition is a critical driver of biogeochemical cycling in forest ecosystems. Among the pathways of deposition, throughfall (TF) - precipitation that has interacted with the forest canopy - plays a significant role in the input of atmospheric substances and canopy leachates to the forest floor. We investigated the influence of airborne pollen deposited in TF on its chemical composition across 60 Level II plots of the ICP Forests network in eight European countries. Based on 196 TF samples collected during the 2018 early vegetative season, we identified 53 pollen taxa, with Pinus, Picea, Fagus, and Quercus accounting for 91.4% of the total grains. Linear mixed-effects models revealed significant positive links between pollen deposition rates and TF fluxes of several ions, especially potassium (K⁺), which was explained by pollen abundance of all four main tree genera. Additionally, pollen of broadleaved taxa (Fagus, Quercus) influenced fluxes of the non-purgeable organic carbon (NPOC), dissolved organic nitrogen (DON), ammonium (NH₄⁺), and phosphate (PO₄^3-). Conversely, broadleaved pollen showed a weak negative link with nitrate (NO₃⁻) fluxes, even if its interpretation requires a cautious approach, as several non-mutually exclusive pathways could explain it. Fractionation of the DOC pools highlighted a higher potential for microbial origin in broadleaves than in conifers. These findings provide the first continental-scale evidence of a link between pollen and TF water chemistry, concurrently analyzed on each water sample. This study underscores the ecological significance of pollen beyond plant reproduction and the need to consider flowering phenology and pollen release in forest deposition and nutrient assessments.