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Abstract
Lower treeline ecotones are influenced by several processes, including fire, grazing, and climatic variables, but rocky substrate is an under-recognized factor affecting their patterns and dynamics. On the Northern Range of Yellowstone National Park, we hypothesized that north-facing slopes and riparian zones would explain the distribution of most forested areas, but that rocky substrate would explain a substantial additional portion of the lower treeline, and that advancement of conifers into grassland has occurred along rocky substrate. We used GIS software and change detection analysis to assess the spatial distribution and change of the forest with respect to north-facing slopes, riparian areas, and rocky substrate. As in other lower treeline studies, we found that north-facing slopes and riparian zones contained the majority of forested land (63%). However, areas with rocky substrate accounted for more forested area than north-facing slopes and riparian zones combined (67.4%), including 28% of forest that was not explained by the first two variables. Change detection analysis between 1954 and 2006 revealed that conifer advancement was insignificant but the number of forested patches increased and the mean size of patches decreased. These results suggest a more fragmented lower treeline environment that may indicate forest advancement in small patches. Field surveys revealed that rocky substrate did not provide conifers with protection from browsing ungulates or fire, yet conifers preferentially established on rocky areas. Our landscape analysis suggests that weathered rock increases soil coarseness in a landscape dominated by fine glacial till, which improves conditions for seedling establishment by reducing competition with grasses and shrubs sufficiently to overcome the negative influence of dry, nutrient-poor microsites in a moisture-stressed environment.