Plant invasions in protected areas at multiple scales:: Linaria vulgaris (Scrophulariaceae) in the west Yellowstone area

Invasive alien plants have long been recognized as a threat to low-elevation, disturbed environments, but the case of Linaria vulgaris Mill. in Yellowstone National Park and Gallatin National Forest shows that invasions can also spread to high-elevation natural reserves. Because invasions in protected areas are a product of complex processes occurring over a broad range of scales, we argue that a multi-scale research approach is needed to capture both patterns and potential mechanisms of the invasion process. Mapping L. vulgaris at the landscape scale, we found the species occupying a broad range of sites, apparently originating from just 2 historical sources, colonizing both human-caused and natural disturbances. Analyzed at the stand scale, patches tend to aggregate in newly invaded areas and disperse in heavily infested areas. The data suggest that patches grow in size by clonal growth and in number by creation of new satellite patches. Radial patch growth rates are related to site characteristics. Clonal patch scale analysis shows that ramet densities and Linaria's effects on native plants are highest in patch centers. Both mean ramet height and reproductive vs. vegetative ramet height ratio are higher in patch cores. These results suggest that L. vulgaris may displace natural vegetation by maintaining vigor even in large and old clonal patches. Our results confirm that L. vulgaris is a significant threat to native biodiversity in open, human- or naturally disturbed environments in protected areas of the Rocky Mountains. A multi-scale method can allow managers to better understand patterns of invasion and prioritize managemerit activities to control invasive alien plants, especially in heterogeneous protected area landscapes.