Shade is the most important factor limiting growth of a woody range expander

The expansion of woody plants into grasslands and old fields is often ascribed to fire suppression and heavy grazing, especially by domestic livestock. However, it is also recognized that nutrient availability and interspecific competition with grasses and other woody plants play a role in certain habitats. I examined potential factors causing range- and niche expansion by the eastern redcedar Juniperus virginiana, the most widespread conifer in the eastern United States, in multifactorial experiments in a greenhouse. Historical records suggest that the eastern redcedar is a pioneer forest species, and may be replaced as the forest increases in tree density due to shading. Another possible factor that affects its distribution may be nutrient availability, which is higher in old fields and other disturbed lands than in undisturbed habitats. In its historic range, eastern redcedars are particularly abundant on limestone outcrops, often termed ‘cedar barrens’. However, the higher abundance on limestone could be due to reduced interspecific competition rather than a preference for high pH substrates. I manipulated shade, fertilization, lime, and interspecific competition with a common dominant tree, the post oak Quercus stellata. In a separate experiment, I manipulated fire and grass competition. I measured growth rates (height and diameter) and above- and belowground biomass at the end of both experiments. I also measured total non-structural carbohydrates and nitrogen in these plants. Shade was the most important factor limiting the growth rates and biomass of eastern redcedars. I also found that there were significant declines in nitrogen and non-structural carbohydrates when shaded. These results are consistent with the notion that the eastern redcedar is a pioneer forest species, and that shade is the reason that these redcedars are replaced by other tree species. In the second experiment, I found that a single fire had a negative effect on young trees. There was no significant effect of competition with grass, perhaps because the competitive effect was shading by grasses and not nutrient depletion. Overall, the effects of shade were far more apparent than the effects of fire.