Tree diversity reduces pine infestation by mistletoe

The pattern that a given tree species suffers less damage when growing with heterospecific neighbors than amongst conspecific plants, i.e. associational resistance, is common for insect herbivores and many fungal pathogens. However, associational resistance to parasitic plants has never been tested in a replicated study. Using paired forest plots, we investigated whether tree diversity triggered associational resistance to a tree parasite, the European mistletoe Viscum album ssp. austriacum, by comparing pure stands of Scots pine (Pinus sylvestris) with mixtures of Scots pine and Maritime pine (Pinus pinaster) in northern Spain. Maritime pine, with 1.2% of trees being infested, was considered a non-host species in the study area. The infestation level of Scots pines was significantly higher in pure plots (45.1%) than in mixed plots of Scots pines and Maritime pines (25.4%). Our study is the first to quantify associational resistance to a plant parasite in mixed vs. pure forest stands and suggests that mechanisms proposed to explain associational resistance to insects and pathogens also apply to plant parasites. Scots pine trees that were taller than the surrounding trees had a higher infestation probability, in both pure and mixed stands. Scots pine trees growing in mixtures were slightly lower than Maritime pines, suggesting that associational resistance was partly driven by reduced relative tree height. However, the effect of plot type (pure vs. mixed) remained significant after the effect of tree height was accounted for, thus indicating that other factors also contributed to lower mistletoe infestation in mixed plots. In particular, the behavior of birds dispersing mistletoe seeds might differ in mixed vs. pure stands.