Are mixtures of aspen and white spruce more productive than single species stands?

In boreal mixedwood forests, aspen (Populus tremuloides) and white spruce (Picea glauca) commonly grow in mixture. These species may avoid competition through differential shade tolerance, physical separation of canopies, phenological differences, successional separation, and differences in soil resource utilization. Aspen may also be able to positively affect the growth of white spruce by improving litter decomposition and nutrient cycling rates, controlling grass and shrub competition, ameliorating environmental extremes, and reducing pest attack. These positive relationships likely make mixed-species stands more productive than pure stands of the same species. The evidence regarding the productivity of pure versus mixed aspen/white spruce stands in natural unmanaged forests is examined in this paper.