The ability of individual animals to move across complex landscapes is critical for maintaining regional populations in the short term (Fahrig 2003; Cushman 2006), and for species to shift their geographic range in response to climate change (Heller & Zavaleta 2009). As organisms move through spatially complex landscapes, they respond to multiple biotic and abiotic factors to maximize access to resources and mates while minimizing fitness costs such as mortality risks. Habitat fragmentation decreases dispersal success (Gibbs 1998), increases mortality (Fahrig et al. 1995) and reduces genetic diversity (Reh & Seitz 1990; Wilson & Provan 2003). Local populations may decline if immigration is prevented (Brown & Kodric-Brown 1977; Harrison 1991) and may prevent recolonization following local extinction (Semlitsch & Bodie 1998).