Do birds see the forest for the trees? Scale-dependent effects of tree diversity on avian predation of artificial larvae

The enemies hypothesis states that reduced insect herbivory in mixed-species stands can be attributed to more effective top-down control by predators with increasing plant diversity. Although evidence for this mechanism exists for invertebrate predators, studies on avian predation are comparatively rare and have not explicitly tested the effects of diversity at different spatial scales, even though heterogeneity at macro- and micro-scales can influence bird foraging selection. We studied bird predation in an established forest diversity experiment in SW Finland, using artificial larvae installed on birch, alder and pine trees. Effects of tree species diversity and densities on bird predation were tested at two different scales: between plots and within the neighbourhood around focal trees. At the neighbourhood scale, birds preferentially foraged on focal trees surrounded by a higher diversity of neighbours. However, predation rates did not increase with tree species richness at the plot level and were instead negatively affected by tree height variation within the plot. The highest probability of predation was observed on pine, and rates of predation increased with the density of pine regardless of scale. Strong tree species preferences observed may be due to a combination of innate bird species preferences and opportunistic foraging on profitable-looking artificial prey. This study therefore finds partial support for the enemies hypothesis and highlights the importance of spatial scale and focal tree species in modifying trophic interactions between avian predators and insect herbivores in forest ecosystems.