Up against the edge: invasive species as testbeds for basic questions about evolution in heterogeneous environments.

Yogi Berra is often credited with having opined that 'prediction is very difficult, especially about the future'. There is no discipline for which this statement holds with more force than invasion biology, where it has been historically very challenging to predict the fate of introduced species (Williamson 2006). Some species after introduction quickly go extinct. Other relatively similar species may persist, but with little spread from their initial beachheads. Yet others can become aggressive invaders, with devastating consequences for native communities and ecosystems. This lack of predictability may of course sometimes reflect a simple lack of knowledge, both about key features of a species' basic biology, and about the environmental and community milieu in which invasion occurs (Williamson 2006). However, unpredictability may also arise from a fundamental fact about populations of living organisms - they almost always contain genetic variation, and so are not fixed entities responding to an environmental template, but instead labile in how they cope with the environment, over many spatial and temporal scales. Chance vicissitudes in the origination, maintenance and spatial organization of genetic variation could play a large role in generating the observed unpredictability in the fates of introduced species. The degree to which a particular introduced species becomes 'invasive'- to the extent of coming to the attention of worried land managers, governmental officials and the public - may reflect in part its capacity for adaptive evolution across a wide range of environmental conditions.